Belaoussoff Et Al. 2003. Assessing Tillage Disturbance On Assemblages of Ground Beetles

Biodiversity and Conservation 12: 851882, 2003.
2003 Kluwer Academic Publishers. Printed in the Netherlands.
Assessing tillage disturbance on assemblages of

ground beetles (Coleoptera: Carabidae) by using a
range of ecological indices
SVENJA BELAOUSSOFF 1 , PETER G. KEVAN 2, *, STEPHEN MURPHY 3 and
CLARENCE SWANTON 4
1
School of Rural Extension Studies, University of Guelph, Guelph, Ontario, Canada N1 G 2 W1;
Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada N1 G 2 W1;
3
Department of Environmental and Resource Studies, University of Waterloo, Waterloo, Ontario,
Canada N2 L 3 G1; 4 Department of Plant Agriculture, University of Guelph, Guelph, Ontario, Canada
N1 G 2 W1; * Author for correspondence (e-mail: pkevan@ uoguelph.ca; fax: 11 -519 -837 -0442)
2
Received 14 June 2001; accepted in revised form 29 April 2002
Key words: Carabid beetles, Disturbance, Diversity and abundance distributions, Diversity indices,
Tillage
Abstract. Many ecological studies have used diversity indices to assess the impact of environmental
disturbance. In particular, ground beetles have been advocated as a good group for assessing disturbance.
Most studies on various organisms have used only one or two indices. For our study of the impact of
tillage disturbance on carabid beetles in farm fields in southern Ontario, Canada, we used seven different
diversity indices (richness, ShannonWiener, BergerParker, Q-statistic, Margalef, a and evenness).
Few studies have used deviation from diversity abundance models as a measure of disturbance; however,
we use three that are applicable to our data (geometric, log-normal and log-series). The indices and
models were used to test the null hypothesis that there is no change in diversity with increasing tillage
disturbance, and that there is no difference in diversity with different crops or years. We were not able to
reject the null hypothesis that there is any diversity difference among farms. We also found that there was
no single diversity index or model that was better than any other at detecting disturbance. These results
are supplemented by a meta-analysis of 45 published data sets for the same taxon but in different habitats.
The meta-analysis supports the conclusions from our field research that diversity indices and models are
not useful for detecting the possible effect of disturbance on assemblages of carabid beetles.
Introduction
Ecologists often use changes in species diversity to determine the effects of
disturbance because it is an important component of any ecosystem (May 1975;
Hutchinson 1978; Magurran 1988). To determine the effect of disturbance on
species diversity it is necessary to use appropriate species assemblages as indicators
of disturbance. Species assemblages above the single species level are communities,
functional groups, guilds and taxocenes (Belaoussoff 2000). In this study we used
functional groups. Functional group can be defined as a group of not necessarily
related species exploiting a common resource base in a similar fashion. Within a
functional group there is greater similarity in ecological resource requirements than
within a guild, thereby implying that there is a greater degree of interspecific
852
competition (Arthur 1984; Colwell and Winkler 1984). Some examples of functional groups include pollinating bees (Kevan et al. 1997), livestock gut fauna (Izsak
and Hunter 1992), and herbivorous Macrolepidoptera (Laroca et al. 1989; Hill et al.
1995).
We chose to use functional group because there is an overlap in resource
requirements between species in a functional group. Disturbances would affect
those species evenly by disrupting resources that they all use. Species within
communities and guilds often depend on different resources. Hence, a given
disturbance would affect some of those resources but not all, and therefore not all
species would be equally affected. Because the species assemblages which make up
communities and guilds are not equally affected by disturbances, they would not be
as sensitive to disturbance as functional groups would be.
There are two different approaches to assess the effect of disturbance on species
within functional groups: differences in diversity indices, and deviation from
diversity and abundance models. Of the two, diversity indices are more frequently
used by ecologists. Diversity indices are generally simple to calculate (with some
exceptions, e.g., the Q-statistic; also see Magurran 1988), especially in comparison
to the mathematics and statistics behind diversity and abundance models. Although
there are many different diversity indices, they generally fall into one of two
categories: species richness indices, and indices based on the proportional abundances of species (Magurran 1988). Richness indices are a measure of the number
of species, or individuals, in a defined sampling unit (e.g., number of species,
density, abundance). Indices which measure the proportional abundance of species
combine species richness and evenness (the component of species diversity that
measures the relative abundance of species) into a single figure (Magurran 1988)
(e.g., ShannonWiener, BergerParker, Q-statistic, Margalef, a and evenness
itself). These indices tend to be biased either towards species richness (e.g., a,
Q-statistic) or dominance (e.g., BergerParker) (Magurran 1988).
Diversity indices can be useful because they provide rapid, and easily calculated,
ecological measures. Also, because many researchers use indices, it is possible to
make some comparisons between similar studies which use the same indices.
Nevertheless, there are three main shortcomings of these indices. First, they can be
used only for comparisons between sites, or on sites over time, or both [for e.g.,
studies on the effect of mining waste on fish (Cornell et al. 1976), landfill
reclamation on invertebrate populations (Judd and Mason 1995) and altitude on
spider populations (Otto and Svennson 1982)]. Second, diversity indices are only
statistical artifacts and do not have any intrinsic biological meaning (Southwood
1978; Magurran 1988). Third, the use of different indices with the same data can
result in different conclusions (Belaoussoff 2000).
One way of overcoming weaknesses in diversity indices is to have a diversity
measure which is stand-alone and does not need inter-site or inter-time comparisons. With such a stand-alone measure, a theoretical standard against which to
make comparisons is required. To avoid the shortcoming of being a statistical
artifice, the standard should be based on community structure, as well as ecological
and evolutionary theory. It is possible to compare the structure of a functional group
853
against the standard and to assess whether or not the functional group is disrupted by
a disturbance. Theoretically, if the structure of the functional group fits the standard,
then it is not disturbed. If it deviates from the standard, then the group is affected by
the disturbance. Diversity and abundance models, and distributions of size within
functional groups, provide a good starting point to find the standards needed for
stand-alone measures of diversity, because they are based in ecological and
evolutionary theory.
Although diversity and abundance models have been present in the ecological
literature for over 80 years (e.g., Raukiaer 1918), diversity indices are more
frequently used by ecologists to examine the effect of disturbance on species
diversity (Southwood 1978; Magurran 1988). Relationships describing the diversity
and relative abundance of species in a community were originally suggested by
Raukiaer (1918), and then again by Motomura (1932). Although different diversity
and abundance models have been developed (e.g., Mandelbrots (1977) Zipf
Mandelbrot model; Dewdneys (1997) logistic J and Hubbles (personal communication) zero-sum multinomial), four principal distributions of diversity and abundance (MacArthurs broken stick model, geometric series, log series and log
normal) remain central to discussions about diversity and abundance models
(Magurran 1988). Because those four distributions are commonly considered, they
are the ones used in this study. Furthermore, some of the alternate models are not
based on ecological principles [e.g., the ZipfMandelbrot model is based on
information theory (Magurran 1988)], or are empirical fits of data which cannot be
separated from log normal distributions [e.g., the logistic J model (Dewdney
1997)].
The broken stick model reflects equally divided niche space (MacArthur 1957).
Its biological meaning is not obvious (Hutchinson 1978; Magurran 1988), although
empirically it is most likely to occur in groups of somewhat mobile animals living in
a homogeneous environment. Unlike the other models, this model reflects an
equitable state (Magurran 1988) and there is no competition between members of
the functional group. The value of the broken stick model has been challenged
(Wilson 1993), and there are few examples of where it occurs [e.g., passerine birds
(MacArthur 1960) and minnows (King 1964)].
Both the geometric and log series distributions are believed to result when niche
pre-emption occurs [i.e., a few species pre-empt, or occupy, most of the niche
(Hutchinson 1978)]. In these distributions few species dominate, and those few
species occupy a large portion of the available niche space. Field data have shown
that geometric distributions are found in species-poor (often harsh) environments
which are in the very early stages of succession (Whittaker 1965, 1972, 1977) or
under disturbed conditions (e.g., Kempton and Taylor 1974). As succession
proceeds, or as conditions improve, species abundance distributions become log
series (Magurran 1988).
The log normal relationship describing the diversity and relative abundance of
species in a community is one of the oldest proposed (Raukiaer 1918; Motomura
1932). In communities described by this distribution there are few common species,
more less common species, and few rare species. It was Preston (1948) who first
854
tested the idea that species diversity and abundance can be described by log
normally distributed frequencies. Since then the model has been shown to fit many
biological communities [e.g., diatoms (Patrick 1968); soil arthropods (Hairston and
Byers 1954); Macrolepidoptera (Kempton and Taylor 1974; Laroca and Mielke
1975; Laroca et al. 1989; Hill et al. 1995), birds and mammals (Preston 1962); bees
(Mackay and Knerer 1979; Tepedino and Stanton 1981; de Bortoli and Laroca 1990;
Barbola I. de and Laroca 1993; Stubblefield et al. 1993; Kevan et al. 1997) and
Salmonella strains in livestock (Izsak and Hunter 1992)].
The log normal model of diversity and abundance is more than a statistical
property of sampling (cf. May 1975). Sugihara (1980) suggested that the niche space
of a taxonomically related group is sequentially split by the constituent species, and
the fraction of the niche space apportioned to a species is proportional to its
abundance. The log normal model, as described by Sugihara, is also known as
Sugiharas sequential breakage model. Until Sugihara (1980) broached its biological rationale, the log normal model was considered to be a statistical consequence of the Central Limit Theorem (MacArthur 1960; May 1975; Tokeshi 1993).
The model represents an organizational pattern for a functional group, and is based
on the following considerations (Sugihara 1980):
1. the existence of an hierarchical niche structure in the functional group;
2. the underlying structure of niches is reflected in the pattern of abundance;
3. it is applicable to functional groups.
A hierarchical niche structure implies that each species making up a functional
group is different from the others and none is redundant. Tilman and Downings
study (1994; see above) supports that idea and links diversity to ecosystem stability.
Because the log normal model of diversity and abundance has biological
meaning, it can be used to address two ecological issues: (1) how to quantitatively
define a functional group, and (2) the need for an indicator of disturbance which is
free from temporal, or spatial, comparisons of species. Generally, researchers
classify species assemblages as functional groups, guilds or taxocenes because the
species within the assemblage are closely related, occur together in a particular
habitat or seem to have similar ecological niches. Competition occurs between
constituent species of functional groups, thus it is possible to assess them quantitatively. It is useful to employ quantitative methods to describe a functional group in
order to give the definition greater scientific validity than if it is merely based on
subjectively grouping species together (Jaksic 1981). By using a fit to the log normal
model under undisturbed conditions it may be possible to assess quantitatively if a
functional group is present. If this way of describing a functional group is not used
with caution, it is possible to place more than one functional group together and still
obtain a log normal distribution of diversity and abundance. Even so, this way of
describing a functional group is an initial, and much needed, quantitative approach
to determining if a collection of species form a functional group. Researchers using
log normality to define functional groups have to be conservative, and cognisant of
the ecological backdrop in which log normality is used.
855
Preston (1980) proposed that the degree of deviation of diversity and abundance
from log normality can be used as an index of the degree of disturbance. The idea
that deviation from log normality might reflect the effect of disturbance has been
tested with diverse assemblages of marine organisms (Gray and Mirza 1979; Gray
1979, 1983). Those studies supported the hypothesis that there would be deviation
from log normality, even though the organisms that were sampled did not belong to
a single functional group. Thousands of individuals were collected, and it is likely
that several functional groups were each well represented and were all similarly
affected by the pollution examined by Gray (1983) and Gray and Mirza (1979). To
date, studies which have associated departure from log normal distributions of
diversity and abundance with disturbance have been published on pollinating bees
(Kevan et al. 1997), Lepidoptera (Laroca et al. 1989; Butler 1992; Butler et al. 1995;
Hill et al. 1995), gut bacteria in livestock (Izsak and Hunter 1992), and soil
arthropods (Hagvar
1994). Using deviation from species diversity and abundance
relationships to determine whether or not a biological community is disturbed is
controversial (Lambshead et al. 1983; Shaw et al. 1983; Nelson 1987). In general,
the major critics of the method (e.g., Nelson 1987) have lumped unrelated, and / or
only partial, functional groups together, thereby dismissing the approach without
understanding why it did not work.
Functional groups exposed to unexpected disturbance events are expected to
deviate from log normality. Such deviation merely raises a red flag, which indicates
that the functional group in question may have been disrupted (Kevan et al. 1997;
Belaoussoff and Kevan 1998). According to some researchers, log normality is a
characteristic of communities in equilibrium (Stenseth 1979; Ugland and Gray
1982). However, a study on birds shows that undisturbed forest habitats are not
necessarily at equilibrium (Gee and Giller 1986). Thus, situations exist in which
functional groups in undisturbed habitats might deviate from expected log normality. Deviation of functional groups away from theoretical standards which are
provided by diversity and abundance is a relatively new approach to studying the
effects of disturbance. As well, deviation of size within functional groups is a novel
approach. More work is needed to sort out the complexities of functional groups and
deviation from log normality as an indicator of disturbance (Hill and Hamer 1998).
Our study examines methods to quantitatively define functional groups with
respect to Carabidae (i.e., carabid or ground beetles) on farms representing a
gradient of tillage disturbance. Carabidae need to be examined as a potential group
for assessing disturbance because they are considered to be sensitive indicators of
environmental change (e.g., Dritschillo and Wanner 1980; Sekulic et al. 1987; Eyre
and Ruston 1989), they are abundant within a wide range of ecosystems (see Thiele
1977), they are relatively easy to catch in large numbers (Thiele 1977), the
taxonomy of many species is well known (Lindroth 19611969) and expert
identification is accessible. Furthermore, more is known about the ecology of
carabids than of almost all other insect families (Thiele 1977; Larochelle 1990).
It is poorly understood how carabids are affected by agricultural disturbance.
Previous studies which have attempted to assess consequences of tillage for carabid
856
diversity were not conclusive. For example, several studies have shown that carabid
diversity is greater in fields under minimum tillage than conventional tillage
(Dritschillo and Wanner 1980; House and All 1981; House and Stinner 1983;
Ferguson and McPherson 1985; Brust et al. 1986; Carcamo

1995; Carcamo
et al.
1995). Other studies either indicate that tillage does not affect carabids (Tyler and
Ellis 1979; Dritschillo and Erwin 1982; Hokkanen and Holopainen 1986; Kromp
1989; Mack and Buckman 1990; Tonhasca 1993), or even that carabids are more
diverse in conventionally tilled fields (Edwards et al. 1979; Barney and Pass 1986).
Some researchers also have found that rotation and crop type affected carabid
diversity (Brust et al. 1986), whereas others (Carcamo

et al. 1995) did not. One
problem with most of these studies is that the only diversity measures that were used
were richness and abundance. Although these measures have intuitive appeal, they
do not provide information about changes in the relative proportions of species, i.e.,
community structure (Magurran 1988). To find out how useful carabid beetles are as
indicator species, it is necessary to fully explore their potential with different
measures of diversity (see above). Although diversity indices have been used with
carabid beetles on other studies, the beetles need to be examined in the context of
deviation from log normal distributions of diversity and abundance.
In this study the disturbance we used was tillage. Tillage is the process of turning
the soil and incorporating organic litter into it, and is an important agricultural
practice. Shifts from high tillage to no tillage practices have been occurring
throughout North America since the 1970s (House and Stinner 1983). The main
types of tillage used in southern Ontario, Canada are: high (mold board plow),
medium (chisel plow) and no (conservation) tillage. In Ontario, no tillage is the least
common of the three practices (since the mid-1990s it has been used on only
1015% of farms). High tillage penetrates approximately twice as far as medium
tillage. No tillage involves seeding with minimal disturbance (Oryokot et al. 1997).
Fields under high and medium tillage, in contrast to those under no tillage, lack
distinguishable soil horizons (Allmaras and Dowdy 1985), have less organic detritus
on top (Allmaras and Dowdy 1985), are much drier (House and Stinner 1983;
Allmaras and Dowdy 1985), and exhibit more soil compaction (House and Stinner
1983; Tonhasca and Stinner 1991). Because increasing tillage decreases the number
of microhabitats available to the beetles and the abundance of their prey (House and
Parmelee 1985; Stinner et al. 1988), carabid diversity is likely to be affected by
tillage. In parallel studies which were conducted on the same sites as the present
study, we found that weed diversity was negatively affected by tillage (Murphy et
al. 2003) while isopod diversity increased with greater degrees of tillage (Belaoussoff et al. 1998).
We used carabid beetles which were collected from four farms that represent a
gradient of tillage disturbance, and then supplemented our findings with a metaanalysis of 45 published data sets. This meta-analysis was an important contribution
to the development of our general conclusion about the usefulness of carabid beetles
as indicator species in disturbance. Our null hypothesis for all diversity indices and
models was that there would be no change with increasing tillage, or other
disturbance.
857
Table 1. Vegetation which was adjacent to all replicates (i.e., fields within farms) used in this study.
Mowed margins
Wood edge
Woods (maplebeech)
Daucus carota
Conyza canadensis
Amaranthus retroflexus
Ambrosia artemisiifolia
Poa pratensis
Phleum pratense
Dactylis glomerata
Bromus inermis
Elytrigia repens
Alliaria petiolata
Acer saccharum
Carpinus caroliniana
Ostrya virginiana
Viburnum acerfolia
Equisetum spp.
Erythronium americanum
Claytonia virginiana
Geranium robertianum
Materials and methods
Selection of agricultural fields

We began with over 100 willing co-operators within the designated area, and visited
over 30 of them. To ensure that any effects of tillage which we might find would
reflect the current tillage regime, we required farms on which the same tillage
practice had been in use for a minimum of 5 years. This constraint was most limiting
for the selection of a no tillage farm, because only 1015% of farmers in the Grand
River Watershed had been using no tillage for at least 5 years before our study
commenced. We wanted to control for as many variables as possible and originally
ended up with three farms which were highly comparable except for tillage practice.
In the second year of the study we found an additional high tillage farm which
neighboured the no tillage farm. All the fields were to be bordered on one side by a
wood lot to assure conformity of edge vegetation on at least one side of the field. We
surveyed the vegetation in the wood lot and field margins surrounding all potential
fields, and we agreed that only fields surrounded by similar vegetation could be used
to assure comparative rigour (Table 1). All of the fields that were used were also
restricted by having more or less the same rectangular shape (approximately twice as
long as wide) and area (approximately 10 ha). The slopes, aspects and soil types of
all fields also had to be similar; however, one of the high tillage farms had slightly
more loamy soil than the other farms (Table 2). Soils were tested as to type using
methods described by Spurr and Barnes (1980).
The average corn heat units for each of the farms used ranged between 2650 and
2750 (Brown and Bootsma 1993). Crops used included corn (Zea mays L. Pioneer
3902, planted in 76 cm rows at 75 000 seeds ha 21 ), soy bean (Glycine max (L.)
Merr. KG 40, planted in 53 cm rows at 50 000 seeds ha 21 ) and winterwheat
(Triticum aestivum L., planted in 18 cm rows at 50 000 seeds ha 21 ). Farmers applied
fertilizer (15, 40, 40 kg ha 21 NPK), except in years following G. max. Glyphosate
was applied at 1.0 kg ai ha 21 as a burn-down herbicide in all rotations and all years.
When planting Z. mays, farmers applied pre-emergence tank mixes of linuron and
metalochlor (1.1 and 1.9 kg ai ha 21 , respectively). For G. max, pre-emergence tank
mixes of metalochlor and imazethapyr (2.0 and 1.0 kg ai ha 21 , respectively) were
858
Table 2. Summary of physical characteristics of the four treatments (farms representing different degrees
of tillage) used in this study.
Tillage type
Location
Treatment
Replicate
Soil type
Slope
Rotation
No till
Oxford
NT
Moldboard
Oxford
HT1
Chisel
Waterloo
MT
Mold board
Wellington
HT2
NT-1
NT-2
NT-3
NT-4
NT-5
HT1-1
HT1-2
HT1-3
MT-1
MT-2
MT-3
MT-4
MT-5
MT-6
HT2-1
HT2-2
HT2-3
HT2-4
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Sandyloam
Loam
Loam
Loam
Loam
0.25
1.50
1.25
1.00
1.25
1.25
0.25
1.25
1.50
0.75
1.50
0.50
0.50
0.25
0.00
0.00
1.75
1.75
3-crop
3-crop
2-crop
3-crop
2-crop
3-crop
3-crop
3-crop
3-crop
3-crop
3-crop
3-crop
3-crop
3-crop
2-crop
2-crop
2-crop
2-crop
HT1 represents the mold board tillage (i.e., high tillage) treatment in Oxford County and HT2 represents
the mold board tillage treatment in Wellington County. MT is the chisel plowed (i.e., medium tillage)
treatment in Waterloo County. NT is the no tillage treatment in Oxford County. Slope is in degrees. Soil
type was determined using the methods of Spurr and Barnes (1980). Three-crop rotation is cornsoy
beanwheat, two-crop rotation is cornsoy bean. Replicate refers to field within farms.
used. In T. aestivum, pre-emergence tank mixes of dicamba, MCPA, and mecoprop

were all applied at 0.5 kg ai ha 21 . The herbicide rates were in accordance with
recommendations in the Ontario Guide to Weed Control (Ontario Weed Committee
1996).
We ended up with four farms (with a total of 18 fields representing three different
farming systems) in Wellington, Waterloo and Oxford Counties, all within 30 km of
one another (Figure 1). With such limited choice, we decided that use of replicate
fields within the similar farms would be our experimental design. The farms used in
this study are conventionally (mold board) tilled, chisel plowed or no till. For the
remainder of this paper, fields within farms which were tilled with a mold board
plow are called HT (i.e., high tillage), the chisel plowed fields are called MT (i.e.,
medium tillage) and the no tillage fields are called NT (i.e., no tillage). In Oxford
county, HT1 is adjacent to NT. Other variables are crop type (winter wheat, soy
bean and corn) and crop rotation (cornsoybean, cornsoybeanwinter wheat). The
HT farm in Oxford County (HT1) is located on Bedford-Blenheim Road, Drumbo.
The NT farm in Oxford County (NT) is located on County Road 22, Drumbo. The
MT farm in Waterloo County (MT) is located on County Road 97, Ayr. In 1995, an
additional high tillage cooperator was included (in Oxford) (HT1) whose farm had
three fields which were within the constraints outlined above, and the HT2
cooperator gave us access to more fields, of which only two additional fields which
were within the constraints outlined above.
859
Figure 1. Locations of the four farms which represent different degrees of tillage in southern Ontario.
HT1 and NT were in Drumbo, Oxford County; HT2 was in Elora, Wellington County; and MT was in
Ayr, Waterloo County. See text for descriptions of the farms.
Pitfall traps for the collection of carabid beetles

Carabids were collected from 1994 to 1997. Each replicate (i.e., field) had a transect
of seven single pitfall traps. The traps consisted of white plastic tubs which were 8
cm deep and 15 cm in diameter. The location of the transect within each replicate
was chosen arbitrarily but began next to the woodlot border, and ran at right angles
to the border. The transect was at least 30 m from the three other edges of the field.
For each transect single traps were placed at the crop edge border, 1, 2, 4, 8, 16 and
32 m into the crop. Additionally, in 1996 and 1997, two barrier traps were put into
place into soy and corn replicates to increase sample sizes. The first trap was placed
1 m from the crop edge nearest the woodlot and the second trap was placed 32 m
from the edge. Barrier traps were not put into wheat replicates, because to do so
would have caused much trampling damage around the trap during installation. The
barrier traps were first tried out in 1996 in NT and HT1 fields, and then put into all
non-wheat replicates in 1997. The two barrier traps were also placed arbitrarily
along a transect in the replicates, but were separated from the single trap transects by
at least 30 m. Barrier traps consisted of 2 m long 3 15 cm high wooden boards
which were fitted into a cross pattern and sunk about 3 cm into the ground. The end
of each board was notched so that a plastic tub could be placed flush against the
860
board when the trap was put into the soil. Bamboo poles of 2.5 m height were placed
beside each trap to indicate its location.
Each year, the traps were installed in early June, after corn and soy crops were
planted and weather permitting. The traps were filled with water and two drops of
detergent to break surface tension. Once a week all the sites were visited and trap
contents were emptied into 70% ethanol for storage until carabid beetles could be
identified. Because trapping efforts were made while crops were on the replicate,
collections were made until mid-August, after wheat was harvested.
Species identification
A key based on Lindroth (19611969) was developed to help with the identification
of the species found in the collection (Belaoussoff 2000). Voucher specimens have
been deposited in the University of Guelph insect museum.
Diversity statistics
The diversity indices which were calculated include richness, the ShannonWiener
index, evenness, the BergerParker index, the a index, Q-statistic, and the Margalef
index (Taylor et al. 1976; Southwood 1978; Magurran 1988). These indices were
calculated for carabid beetle assemblages collected from replicates in HT1, HT2,
MT and NT. Jaccards similarity index was also calculated to determine within-year
similarities between and within replicates in the treatments.
ShannonWiener Index (H9) was calculated as:
Os p lnp d
H952
(1)
where pi 5 Ni /N. Ni is the number of individuals of the ith species in the sample and
N is the total number of individuals.
Shannon evenness was then calculated using the formula:
E5H9 / lnS
(2)
where H9 is the ShannonWiener index and S is the number of species.

The Margalef index (D) was calculated as:
D5sS21d / lnN
(3)
where S is the number of species and N is the number of individuals.

The BergerParker dominance index (d) was calculated from the equation:
d5Nmax /Nt
(4)
where Nmax is the number of the most abundant species and Nt is the total number of
species. In order to ensure that the index increases with increasing diversity, the
reciprocal form of the measure was used (Magurran 1988).
The Q-statistic (Q), which is a measure of the inter-quartile slope of the
cumulative species abundance curve, was calculated first by choosing the upper and
861
lower quartiles (i.e., the points below which 25 and 75% of the species abundances
lie) (Kempton and Taylor 1974). Once the quartiles are selected, the Q-statistic was
calculated by:
On 10.5n
Q5 0.5n R 1 1
R2
D / logsR
/R 1d
(5)
where 0.5n R 1 is half the number of species in the class in which the lower quartile
falls, on r is the total number of species between quartiles, 0.5n R 2 is half the number
of species in the class in which the upper quartile falls, and R 1 and R 2 are the
number of individuals in the classes with the lower and upper quartiles, respectively.
The a statistic (a ) was calculated by:
a5Ns12xd /x
(6)
where x is determined from S /N 5 [(12x) /x][2ln(12x)], S is the total number of

species and N is the total number of individuals.
The above diversity indices are measures of a diversity, which is the diversity of
a biological community within a habitat (Southwood 1978; Magurran 1988). b
diversity is a comparison of species composition between different communities
(Southwood 1978; Magurran 1988). To assess b diversity, the Jaccard index (J)
was used (Southwood 1978):
J5Sab /sSa 1Sb 2Sbd
(7)
where Sab represents the number of species shared by community a and community
b, and Sa and Sb are the number of species found in community a and b, respectively.
Dendrograms based on the Sorensen quantitative index were prepared for carabid
assemblages between and within treatments (Magurran 1988).
Statistical analysis
Data were collected on the 18 fields (replicates) within four treatments (farms). The
treatments represented three different tillage types (no tillage, medium, or high).
Data from the two high tillage treatments consisted of two farms which were treated
separately because one of the farms was added a year into the experiment. Each
replicate was subject to a particular trap type (single or barrier) and crop (corn, soy,
or wheat), where the crop type rotated year to year. Data were examined and the
data were found to be approximately normally distributed. Repeated measures
analysis (RMA) of seven diversity indices (see above) were taken weekly for 9
weeks on each replicate at each position along a transect over the course of 4 years
(19941997). The SAS procedure PROC MIXED (Littell et al. 1996) was used to fit
a mixed model where the effect of replicates within treatments was treated as a
random effect, while the other factors, treatment, crop, trap type, and year remained
fixed. All two-way interactions were included as well. F tests (Littell et al. 1996)
were used to test the overall effect of the fixed factors (treatments), while individual
t-tests were used to reveal differences in the least square means of the levels of each
treatment.
862
In addition to examining diversity index differences between treatments, RMA
was performed on carabid abundance and richness data collected from each replicate
over the 9-week sampling period in each year (time). This analysis was done to
examine if there were differences in abundance, or richness, within a given year.
First, a linear regression of abundance (or richness) over time was fitted to generate a
slope for each treatment combination (treatment, crop, trap type). These slopes were
subsequently analyzed by fitting the model explained above. This analysis was
repeated using the slopes from regressing abundance (or richness) on the position of
the traps along the transect in order to examine if there were trends in the abundance
or richness over the different positions of the traps within a given year.
To compare the results of our study with those on other carabid populations,
published data sets from various disturbance studies were gathered for meta
analysis. Disturbances in those studies include: clear cutting (Sustek
1981; Lenski
1983; Hejkal
1982), pollution exposure (Critchley 1972; Freitag et al. 1973; Jarosik
1985; Asteraki et al. 1992), forest fragmentation (Niemela et al. 1992; Halme and
Niemela 1993) and agriculture (House and All 1981). The same species diversity
indices as used for carabids in this study were applied to those data.
Diversity and abundance models
Carabid data from each treatment were pooled to ensure that the sample sizes would
be sufficient to allow for statistically rigorous testing of differences between
diversity and abundance models. Three diversity and abundance models were
examined: geometric, log series and log normal. The broken stick model was not
tested for, because the data rank abundance graphs indicated that the data did not
conform to that model.
Magurran (1988) suggests that the simplest method to assess whether or not a
geometric series represents a biological community is to use rank abundance curves.
The log series distribution is represented by:
ax,sax 2d / 2,sax 3d / 3, . . . sax nd /n
(8)
2
a x is the number of species predicted to have one individual, (a x ) / 2 those with

two and so forth (Fisher et al. 1943; Poole 1974; May 1975). Magurran (1988)
outlines the procedure to obtain a as follows. The total number of species, S, is
obtained by adding all the terms in the series, which reduces to:
S5a[2lns12xd]
(9)
with x being estimated from:

S /N5s12xd /x[2lns12xdj
(10)
where N is the total number of individuals.

Because a and N summarize the distribution and are related by:
N5alns11N / ad,
the following equation can be used to obtain a :
(11)
863
a5[Ns12xd] /x
(12)
To determine if our treatment data sets conformed to log normal distributions,

they were graduated to a truncated curve using the following general equation
(Raukiaer 1918; Preston 1948; Whittaker 1972):
SsRd5S0 exps2a 2 R 2d
(13)
Species abundances were transformed using log 2 to generate intervals of octaves. S0

is the number of species within the modal abundance octave R 0 . In Equation 13,
S( R) is the number of species in an octave R octaves from R 0 and 1 /a is the width of
]
the distribution a 5 1 /(2s ). This constant, a, for a canonical log normal
relationship has been found to be approximately 0.2 (Hutchinson 1953; Whittaker
1972; Colinvaux 1973).
For species distributions that had modes, log normality was tested. A crude mode
in log normal distributions of diversity and abundance can be seen in histograms as
the value which occurs most frequently but has lower values on either side. Log
normal histograms have modes, but log series graphs do not because the most
frequent value in those histograms is next to the y-axis, and thus has a lower
neighbouring value only on one side. The Lilliefors test, instead of the KolmogorovSmirnov x 2 test, was used to indicate whether or not our data was log
normal because it is more conservative in the probability of rejecting the null
hypothesis (Sokal and Rohlf 1995). This statistical test was recommended by an
anonymous reviewer of Kevan et al. (1997), and also by W. Matthes-Sears
(University of Guelph) and Dr M. Camp (USDA, Maryland), as the most appropriate way to test for fit to log normal distributions. To detect log series distributions, the KolmogorovSmirnov x 2 test was used. As well, rank abundance
graphs and frequency distribution histograms were prepared for pooled carabid data
from each treatment.
Rank abundance distributions were examined to determine if the findings in our
study were similar to those from others [clear cutting (Lenski 1982; Stubblefield et
1983;
al. 1993), pollution exposure (Critchley 1972; Freitag et al. 1973; Jarosik
Hejkal 1985; Asteraki et al. 1992), forest fragmentation (Niemela et al. 1992; Halme
and Niemela 1993) and agriculture (House and All 1981)].
Results
General findings
Thirty genera, and 102 species, of carabids were identified from the 20 495 carabid
beetles collected during this study. Over four years of collection (19941997) 68
species were found on no tillage treatment (NT), 57 on the chisel plowed treatment
(MT), and 42 and 53 on the mold board treatment (HT1 and HT2, respectively)
(Appendix 1). Species which were found on all four treatments include Anisodactylus sanctaecrusis Fabricius, Bembidion inaequale Say, Bembidion tetracolum
864
Say, Bembidion quadrimaculatum oppositum Say, Chlaenius tricolor Dejean,
Harpalus herbivagus Say, Pterostichus melanarius Illiger,
Clivina fossor Linne,
Stenolophus comma Fabricius and Tachys incurvus Say. Agonum muelleri Herbst
occurred frequently on all but NT replicates. Amara Bonelli species were not
collected from HT1, but were found on the other treatments. Cicindela and
Harpalus species were far more common on NT than the other treatments.
Dyschirus species were not found on NT or the neighbouring HT1, but were
collected in low numbers from HT2 and MT replicates.
There were yearly differences in the ranked order of the most numerous species
within each treatment, but the common species within each treatment were present
throughout the study. For HT1 replicates, species which were common throughout
the study period were Clivina fossor, Bembidion quadrimaculatum oppositum,
Pterostichus melanarius, and Tachys incurvus. For HT2 replicates, the most
common species include Agonum muelleri, Pterostichus melanarius, and Bembidion
quadrimaculatum oppositum. For MT replicates, the most common species were
Pterostichus melanarius, and Bembidion quadrimaculatum oppositum. For NT
replicates, the most common species were Harpalus pensylvanicus de Geer,
Pterostichus melanarius, Bembidion quadrimaculatum oppositum and Cicindela
formosa generosa Dejean.
Exotic species which were collected from all treatments were: Agonum muelleri,
Amara aenea, Bembidion tetracolum, Bembidion quadrimaculatum oppositum,
Clivina fossor, Harpalus affinis, Harpalus puncticeps, Pterostichus melanarius,
Trechus discus and Trechus quadristriatus (see Spence (1990) for a list of exotic
species in Canada). The exotic species Bembidion obtusum Serv. was found only on
HT1 and HT2, and Amara familiaris Dft. was collected only from NT. The total
percent abundances of exotic carabids collected from each treatment are: 42.31% in
HT1, 77.48% in HT2, 60.59% in MT and 35.87% in NT. The high percentage
abundance is because some of the most commonly collected species (i.e., Agonum
muelleri, Bembidion quadrimaculatum, Clivina fossor, Pterostichus melanarius) are
exotics. The ratio of exotic species to native species collected from each treatment
was 9:33 in HT1, 10:43 in HT2, 8:49 in MT and 10:58 in NT. The null hypothesis
that exotic species are more common in treatments with less tillage cannot be
rejected when contingency tables are used ( x 2 5 0.38, df 5 3, P . 0.03).
Carabid richness and abundance over transects
For single trap transects, the results of RMA on carabid abundance reveal that there
were no significant effects of farm, year or crop type per trap, for trap positions or
time (F values ranged from 0.56 to 1.84, df ranged from 13 to 1012). For carabid
richness over time there was no overall significant effect for farms (F 5 2.33, df 5
3, 12, P . 0.12); however, there was a significant year by crop interaction (F 5
9.03, df 5 2, 10, P ,, 0.01). Least square means reveal that wheat had a
significantly different effect than did corn and soy (t 5 3.03, df 5 10, P , 0.01, and
t 5 4.17, df 5 10, P ,, 0.01, respectively), but that corn and soy did not differ from
one another.
865
For barrier traps, the results of RMA on carabid richness revealed that there was
no significant effect of treatment or year for trap positions or time (F values ranged
from 0.07 to 2.82, df ranged from 13 to 193). Crop type could not be tested
because barrier traps were only placed in corn and soy fields, and if crop type was
included as a factor the model would not converge. Although for carabid abundance,
there was no overall significant effect of trap position for either treatment or crop (F
values ranged from 0.29 to 0.85, df ranged from 13 and 15), there was a
significant year by time (i.e., collection period) interaction for abundance. Least
square means revealed that NT replicates were significantly different from the other
three treatments in 1997, when more beetles were trapped in barrier traps in NT than
with the other treatments.
Generally, there was no difference in trap position or over collection time for
richness or abundance of carabids collected in fields of the four treatments.
Therefore, it is possible to pool data from the traps over time to calculate diversity
indices for each treatment.
Species diversity
Jaccard s similarity index
Figure 2 and Table 3 provide Jaccards similarity indices for all replicates. The
mean similarity within each treatment is: 0.37 for HT1, 0.46 for HT2, 0.31 for MT
and 0.35 for NT. There is high similarity between transect types (i.e. single and
barrier) within replicates, with values ranging from 0.70 to 0.80. More carabids were
trapped in transects with barrier traps than in transects with single traps. However,
species compositions found in the two trap types were not significantly different.
Dendrograms showed that species composition in NT and MT are more similar to
one another than in HT1 and HT2. Within treatments, species tend to be most similar
between years, but crop type does not seem to have an effect.
Diversity indices
The overall effect of treatment, crop, or transect, on diversity indices (Q-statistic,
Margalef, richness, evenness, ShannonWiener, a and BergerParker) was not
significant. There were some year by treatment interactions, thus it is necessary to
look at the simple effects of treatment by year comparisons with least squares
means. For the Q-statistic, in 1994, HT2 was significantly different from MT and
NT (t 5 2.85, df 5 26, P , 0.01; t 5 2.10, df 5 26, P , 0.05, respectively), and, in
1996, HT1 was significantly different from NT (t 5 2.11, df 5 26, P , 0.04). For
richness, in 1996, NT was significantly different from HT1 and MT (t 5 2.14, df 5
26, P , 0.04; t 5 2.55, df 5 26, P , 0.01, respectively) and, in 1997, NT was
significantly different from HT2 and MT (t 5 2.74, df 5 26, P , 0.01; t 5 2.28, df
5 26, P , 0.03, respectively). For ShannonWiener, in 1994, HT2 was significantly different from NT (t 5 4.56, df 5 26, P ,, 0.01); in 1996, HT1 was
significantly different from MT and NT (t 5 2.27, df 5 26, P , 0.03; t 5 2.28, df 5
26, P , 0.03, respectively). For BergerParker, in 1995, NT was significantly
different from HT2 (t 5 4.12, df 5 26, P ,, 0.03); in 1996, HT1 was significantly
866
Figure 2. Dendrograms of carabid assemblages based on the BrayCurtis index of percent similarity.
HT1 and HT2 represent mold board tillage (i.e., high tillage) treatment in Oxford County and Wellington
County, respectively. MT is the chisel plowed (i.e., medium tillage) treatment in Waterloo County. NT is
the no tillage treatment in Oxford County. The number after the treatment name refers to replicate field. w
5 wheat, s 5 soy, c 5 corn, 94 5 1994, 95 5 1995, 96 5 1996, 97 5 1997.
867
Table 3. Jaccards index of similarity. HT1 and HT2 represent the mold board tillage (i.e., high tillage)
treatment in Oxford County and Wellington County, respectively. MT is the chisel plowed (i.e., medium
tillage) treatment in Waterloo County. NT is the no tillage treatment in Oxford County. The number after
the treatment name refers to replicate field. (a) Between treatments in 1994; (b) between treatments in
1995; (c) between treatments in 1996; (d) between treatments in 1997.
a
NT3 MT1 MT2
HT1-1 0.21 0.20 0.33
NT3
0.21 0.18
MT1
0.21
b
HT2-2 HT2-3 HT1-1 HT1-2 HT1-3 NT2 NT5 MT3 MT4
HT2-1 0.59 0.59 0.40 0.44 0.35 0.13 0.08 0.37 0.35
HT2-2
0.63 0.42 0.47 0.22 0.13 0.14 0.47 0.47
HT2-3
0.42 0.40 0.29 0.13 0.14 0.39 0.38
HT1-1
0.44 0.44 0.13 0.18 0.18 0.44
HT1-2
0.39 0.18 0.11 0.50 0.50
HT1-3
0.18 0.17 0.17 0.29
NT2
0.23 0.07 0.18
NT5
0.09 0.11
MT5
0.40
c
HT2-2 HT2-3 HT1-1 HT1-2 HT1-3 NT1 NT2 NT3 NT4 NT5 MT1 MT2 MT3 MT4 MT5 MT6
HT2-1 0.38 0.32 0.26 0.32 0.28 0.23 0.16 0.33 0.21 0.20 0.30 0.25 0.29 0.31 0.30 0.28
HT2-2
0.54 0.41 0.44 0.25 0.26 0.23 0.42 0.28 0.23 0.38 0.33 0.26 0.40 0.38 0.36
HT2-3
0.35 0.29 0.31 0.23 0.24 0.24 0.32 0.21 0.37 0.24 0.22 0.33 0.37 0.27
HT1-1
0.63 0.39 0.26 0.31 0.29 0.39 0.23 0.27 0.26 0.33 0.53 0.40 0.38
HT1-2
0.30 0.26 0.27 0.23 0.32 0.19 0.27 0.39 0.26 0.33 0.27 0.42
HT1-3
0.14 0.19 0.24 0.21 0.29 0.29 0.30 0.35 0.83 0.35 0.40
NT1
0.48 0.30 0.47 0.29 0.29 0.17 0.14 0.24 0.24 0.19
NT2
0.26 0.60 0.47 0.29 0.18 0.20 0.25 0.25 0.21
NT3
0.33 0.22 0.20 0.15 0.18 0.36 0.20 0.19
NT4
0.40 0.36 0.23 0.17 0.32 0.31 0.26
NT5
0.31 0.22 0.25 0.26 0.31 0.30
MT1
0.32 0.24 0.39 0.43 0.41
MT2
0.25 0.33 0.32 0.36
MT3
0.31 0.44 0.42
MT4
0.39 0.37
MT5
0.63
d
HT2-2 HT2-3 HT1-1 HT1-3 NT1 NT2 NT3 NT4 NT5 MT4 MT5 MT6
HT2-1 0.43 0.39 0.13 0.39 0.26 0.30 0.24 0.17 0.25 0.33 0.33 0.51
HT2-2
0.72 0.11 0.40 0.14 0.19 0.22 0.17 0.18 0.46 0.41 0.37
HT2-3
0.12 0.47 0.18 0.22 0.33 0.23 0.25 0.43 0.35 0.38
HT1-1
0.21 0.07 0.06 0.24 0.08 0.07 0.14 0.14 0.16
HT1-3
0.21 0.22 0.41 0.40 0.46 0.60 0.50 0.49
NT1
0.56 0.39 0.39 0.34 0.33 0.36 0.24
NT2
0.44 0.48 0.44 0.32 0.33 0.28
NT3
0.35 0.55 0.30 0.29 0.36
NT4
0.41 0.32 0.28 0.24
NT5
0.29 0.33 0.36
MT4
0.64 0.41
MT5
0.41
868
different from MT (t 5 3.32, df 5 26, P ,, 0.03) and, in 1997, NT was significantly
different from MT (t 5 2.89, df 5 26, P , 0.01). For a, in 1994, HT2 was
significantly different from MT (t 5 2.89, df 5 26, P , 0.01); in 1996, HT1 was
significantly different from NT (t 5 3.22, df 5 26, P ,, 0.03) and, in 1997, HT1
was significantly different from MT (t 5 2.28, df 5 26, P , 0.03).
Diversity meta-analysis of published data sets on structure of carabid
assemblages
Carabid species diversity was calculated for 45 published carabid data sets from 10
studies (Table 4). As with the carabid data collected for this study, no one diversity
measure was better than the others at pinpointing the sites that were disturbed. For
example, ShannonWiener (H9) measures for Critchleys (1972) data indicated that
the site with greatest exposure to pesticide spraying had greater diversity than the
control site. Meanwhile, Margalefs index suggested the opposite to be true.
Although Lenskis data show that H9 was greater in forested areas than in clear cuts,
Margalef, Q-statistic and a did not.
Species abundance distributions
The species abundance distributions for each farm had long tails of rare species.
This is shown in rank abundance graphs (Figure 3) and in frequency distribution
histograms (Figure 4) of pooled carabid species data for each treatment. The carabid
data did not show a mode in the frequency distribution histograms, and thus the veil
line could not be estimated. Because there is no mode, the data did not conform to
log normality. All the treatments did not conform statistically to log series distributions: for HT1 x 2 5 18.75, df 5 9, P , 0.05; for HT2 x 2 5 23.30, df 5 10, P
, 0.01; for MT x 2 5 21.95, df 5 9, P , 0.05; and for NT x 2 5 33.02, df 5 10,
P ,, 0.01. The high number of rare species, and the straight line suggested in the
rank abundance graphs, suggest that all the distribution abundance patterns for the
carabid communities collected are geometric (Magurran 1988). As with the carabid
populations in this study, all the published carabid species abundance distributions
had long tails of rare species. Because there is no mode, the data did not conform to
log normality. Table 5 lists the log-series analyses for 45 data sets, of which nine
were not log series. In all non-log series data sets there were many rare species,
suggesting that they are geometrically distributed (Magurran 1988).
Discussion
Our study joins the studies which suggest that carabid beetle diversity is not affected
by tillage. Unlike those studies, which mainly used only ShannonWiener and
species richness, we examined a range of different diversity indices, each with
different biases and methods of calculation. We wanted to determine if with less
tillage there is decreased dominance (BergerParker), increased evenness (Shannon
evenness) or richness (a, Q-statistic, species richness, Margalef, ShannonWiener).
869
Table 4. Diversity indices for published carabid data.

Reference
Niemela et al. (1992)
Site
Upland aspen forest

Aspen poplar forest
Spruce bog
Lakeside forest
Meadow
Halme and Niemela (1993) Contiguous forest
Large fragment
Medium fragment
Small fragment
Edge
Surroundings
Sustek
(1981)
Forest
Clearing
Fisher et al. (1943)
Station 1
Station 2
Station 3
Station 4
Station 5
Control
Critchley (1972)
Control
Low dose
High dose
Lenski (1982)
Rms forest
Rms clearcut
Er forest
Er clearcut
Rmn forest
Rmn clearcut
(1983)
Jarosik
Cernpoll
Cernunpoll
Libipoll
Libiunpoll
House and All (1981)
Conventional soy
Conservation soy
Fescue
Field
Woods
Hejkal (1985)
1 month
2 years
6 years
17 years
22 years
Asteraki et al. (1992)
Control 2
After treatment b2
After treatment r2
BP
Al
18
21
13
23
34
16
19
15
25
32
54
18
18
18
14
17
20
17
18
23
23
14
26
31
23
25
19
18
30
28
35
30
19
22
21
21
19
16
22
20
21
18
21
17
12
734
403
342
378
454
437
194
269
405
684
1720
2223
1593
242
302
598
534
984
700
524
582
114
1511
1161
301
499
714
532
2028
1371
3872
2710
2003
3081
3240
7213
1622
347
416
1294
150
75
481
549
651
1.39
1.96
1.81
2.32
2.33
2.18
2.03
1.85
2.30
2.39
2.77
1.88
1.93
2.43
1.99
2.20
2.32
2.04
1.83
1.86
1.93
1.91
2.01
2.54
2.57
2.49
2.18
2.09
1.63
2.09
1.85
2.18
1.70
1.21
1.25
0.93
0.58
1.63
2.22
2.15
2.63
2.09
1.64
1.64
1.51
0.48
0.64
0.71
0.74
0.66
0.79
0.69
0.68
0.71
0.69
0.69
0.65
0.67
0.84
0.75
0.78
0.77
0.72
0.63
0.59
0.62
0.72
0.62
0.74
0.82
0.77
0.74
0.72
0.48
0.63
0.52
0.64
0.58
0.39
0.41
0.31
0.20
0.59
0.72
0.72
0.86
0.72
0.54
0.58
0.61
0.57
0.35
0.33
0.30
0.33
0.23
0.26
0.24
0.27
0.23
0.27
0.41
0.41
0.24
0.29
0.25
0.27
0.37
0.37
0.50
0.46
0.39
0.40
0.32
0.23
0.21
0.27
0.29
0.56
0.35
0.58
0.26
0.48
0.69
0.67
0.79
0.72
0.40
0.35
0.22
0.17
0.35
0.50
0.46
0.49
2.58
3.33
2.06
3.71
5.39
2.47
3.42
2.50
4.00
4.75
7.11
2.21
2.31
3.10
2.28
2.50
3.03
2.32
2.59
3.51
3.46
2.74
3.42
4.25
3.85
3.86
2.74
2.71
3.81
3.74
4.12
3.67
2.37
2.61
2.47
2.25
2.44
2.56
3.48
2.65
3.99
3.94
3.24
2.54
1.70
6.92
19.90
5.30
13.30
12.87
6.92
9.48
5.31
11.01
15.43
23.17
6.38
7.71
10.06
9.56
8.48
8.45
12.16
18.79
19.93
12.50
11.53
14.95
16.14
12.58
12.49
13.21
9.60
14.50
11.50
13.59
10.60
5.56
9.51
9.86
6.52
8.00
14.54
9.97
6.39
11.52
14.11
24.10
10.92
6.21
3.34
4.42
4.64
5.39
8.51
3.26
5.22
3.43
5.89
6.96
10.35
2.67
2.84
4.47
3.04
3.26
4.10
2.92
3.37
4.92
4.78
4.19
4.40
5.85
6.07
5.54
3.59
3.60
4.99
4.98
93.38
4.72
2.91
3.20
3.01
2.66
3.40
4.94
3.48
4.95
3.36
0.34
4.48
3.33
2.96
R is richness, A is abundance, H is the ShannonWiener index, E is the Shannon evenness index, BP is

the BergerParker index, Al is the a diversity index, M is the Margalef index, Q is the Q-statistic. For
descriptions of the sites see the respective studies.
870
Figure 3. Rank abundance plots of carabid beetles collected in our study on a logarithmic scale against
the species rank, in order from the most abundant to the least abundant species. HT1 and HT2 are high
tillage treatments, MT is the medium tillage treatment, NT is the no tillage treatment.
Figure 4. Frequency distributions of carabid species collected from treatments representing high tillage
(HT1 and HT2), medium tillage (MT) and no tillage (NT), in relation to their abundance.
Even though we examined a range of diversity indices, we found that none of the
indexes provided a consistent indication that carabid diversity was different between
crops or farms. These results are supported by our meta-analysis of published
carabid data sets, in which none of the different diversity indices could predict
871
Table 5. Summary of x 2 analysis for log series distributions for data sets in our meta-analysis.
Reference
Site
Log-series
Sig
Niemela et al. (1992)
Upland aspen forest

Aspen poplar forest
Spruce bog
Lakeside forest
Meadow
Contiguous forest
Large fragment
Medium fragment
Small fragment
Edge
Surroundings
Forest
Clearing
Station 1
Station 2
Station 3
Station 4
Station 5
Control
Control
Low dose
High dose
Rms forest
Rms clearcut
Er forest
Er clearcut
Rmn forest
Rmn clearcut
Cernpoll
Cernunpoll
Libipoll
Libiunpoll
Conventional soy
Conservation soy
Fescue
Field
Woods
1 month
2 years
6 years
17 years
22 years
Control 2
After treatment b2
After treatment r2
18
21
13
23
34
16
19
15
25
32
54
18
18
18
14
17
20
17
18
23
23
14
26
31
23
25
19
18
30
28
35
30
19
22
21
21
19
16
22
20
21
18
21
17
12
734
403
342
378
454
437
194
269
405
684
1720
2223
1593
242
302
598
534
984
700
524
582
114
1511
1161
301
499
714
532
2028
1371
3872
2710
2003
3081
3240
7213
1622
347
416
1294
150
75
481
549
651
12.95
7.88
9.16
1.48
36.34
1724.00
11.42
5.21
14.36
11.11
14.31
8.10
9.91
5.96
9.50
5.93
3.49
5.28
16.42
12.15
19.96
7.40
12.91
12.11
2.33
3.27
19.11
14.01
8.92
7.91
3.53
6.17
6.82
9.54
11.88
6.74
7.60
11.65
7.37
13.51
14.33
25.89
13.04
5.78
3.42
0.10 . P . 0.05
P . 0.05
0.10 . P . 0.05
P . 0.95
P , 0.001
P . 0.75
P , 0.05
P . 0.75
0.05 . P . 0.025
0.10 . P . 0.05
0.05 . P . 0.025
P . 0.25
P . 0.25
P . 0.25
0.10 . P . 0.05
P . 0.50
P . 0.75
P . 0.50
0.025 . P . 0.01
0.10 . P . 0.05
0.025 . P . 0.01
0.025 . P . 0.01
0.10 . P . 0.05
P . 0.25
P . 0.75
P . 0.75
0.01 . P . 0.005
0.10 . P . 0.05
P . 0.25
P . 0.25
P . 0.75
P . 0.50
P . 0.25
P . 0.50
P . 0.25
P . 0.75
P . 0.75
0.25 . P . 0.10
P . 0.25
0.10 . P . 0.05
0.05 . P . 0.025
P ,, 0.001
0.10 . P . 0.05
P . 0.50
P . 0.75
Halme and Niemela (1993)
Sustek
(1981)
Fisher et al. (1943)
Critchley (1972)
Lenski (1982)
(1983)
Jarosik
House and All (1981)
Hejkal (1985)
Asteraki et al. (1992)
R is richness and A is abundance. For descriptions of the sites see the respective studies.
872
disturbance any more reliably than any of the others. This suggests that diversity
indices do not provide reliable tools for detecting disturbance when using assemblages of carabid beetles as bioindicators.
Indices are commonly used because ecologists consider them as a standard
approach to addressing questions of disturbance. Just because they are commonly
used does not, however, mean that they are reliable. The indices which are used vary
from study to study, and perhaps the results which researchers use are the ones with
which the null hypothesis can be rejected (Peters 1991). Because we found that
indices are not reliable measures of disturbance, we tried to use diversity abundance
models to describe disturbance in our study.
We found that the diversity and abundance of carabid beetle assemblages in our
study (Figures 3 and 4), and in published studies (Table 5), are all either log series
or geometrically distributed. Log normal distribution of diversity and abundance is
not represented in any carabid assemblage, even in undisturbed sites; thus, carabid
beetles cannot be used as indicators of disturbance when deviation from log normal
distribution of diversity and abundance is used. These findings can be explained in
part by carabid beetle sizes.
Because of large size differences among species, it is questionable whether
competition occurs between many carabid species. Carabid feeding behaviour is
body size dependent (Muller

1985), and between carabid species, size similarity is
an indirect measure of likeness in prey preference (den Boer 1980; Brust and House
1988). The more than 10-fold size difference from the smallest to the largest beetle
species strongly suggests that they consume different foods and that they occupy
separate portions of the micro habitat (Thiele 1977). Maximum relative prey size
tends to increase with the body size. Small species (less than 10 mm long) feed only
on prey with smaller body sizes (Loreau 1988) because they are limited by their
ability to catch, or chew, a number of prey types (Hengeveld 1980). In comparison,
large carabid species (longer than 10 mm) are not restricted to smaller prey (Loreau
1988). As a result, large carabid species tend to have a wider prey spectrum than do
smaller species (Rushton et al. 1991). Additionally, large species generally use a
disproportionately wider share of the resources within local ecosystems (Brown and
Maurer 1986; Siemann et al. 1996). Thus, although data for Carabidae are wanting,
it is reasonable to suggest that larger carabid species catch more, and a greater
diversity of, prey.
The species assemblages which should be used when measuring disturbance by
examining the degree of deviation away from log normality are functional groups.
Carabid beetles, it appears, are not useful as such indicators of disturbance, because
they are not a single functional group. There is likely to be little interspecific
competition between carabids because of their size differences, but also because
carabid feeding behavior is diverse (Larochelle 1990). With the exception of a few
specialists, most carabid species are opportunistic feeders even though they are
often considered as predators (Thiele 1977). They consume what is available and
are not necessarily limited by availability of specific foods (Larochelle 1990).
Further, not only are feeding habits of adults diverse, between larvae there are
873
additional differences in feeding habits (Lindroth 19611969; Thiele 1977). As a
result of the diversity of feeding behaviours during different life stages of the
beetles, disturbances are not expected to have a uniform effect on all species.
Because adults and larvae have different feeding behaviours, there is not necessarily
direct competition between life stages, which in turn suggests that there may not be
competition between all of the carabid species.
Since carabid beetles as a group have feeding behaviors which depend on species
size and feeding preference, they cannot be considered as a single functional group.
Rather, they should be splintered into several size determined groups (i.e., small,
medium and large) so that there would be increased likelihood that competition
occurs within a designated group. To become a complete functional group, each of
those sub-groups must be united with additional species from other taxa. For
example, carabid beetles together with staphylinid beetles of similar sizes and
feeding behaviours may make a functional group which can be analyzed for log
normality. Unfortunately this idea is very difficult to test, because staphylinid
beetles are limited by taxonomic factors and the ecology of many of the species is
not well documented.
Log series and geometric distributions are usually considered to result from niche
pre-emption (Boswell and Patil 1971; May 1975; Thomas and Shattock 1986;
Magurran 1988); however, for carabids this argument cannot be invoked. If nichepreemption occurred, the size of the species which dominate from one year to the
next would be consistent. In this study, and in others, the size of carabid species
which dominate from year to year is not consistent (Belaoussoff 2000). If the
carabid assemblage is made up of a number of incomplete functional groups, then a
log series distribution is to be expected. Within each size limited sub-group there
would be changes in dominance, and carabids would not necessarily be the taxon to
which the dominant species would belong. By haphazardly categorizing members
from different incomplete functional groups together, a log series, or geometric,
distribution would be expected.
Carabids are not effective as indicators for a detailed understanding of the
dynamics of disturbance within ecosystems, because they are not a functional group.
These findings that carabid beetles are not useful as disturbance indicators are
important, because they lead to hypotheses about the types of organisms which are
useful. Communities which should be targeted for providing potential standards
based on diversity and abundance models should include species which feed on a
single type of food, species which do not have size dependent feeding, and species
which are more or less a complete functional group.
874
Appendix 1
Percent abundance of all carabid species collected from various treatments.
Species
High tillage treatment HT1 from 19941997
Tachys incurvus
Pterostichus melanarius
Clivina fossor
Pterostichus lucublandus
Bembidion quadrimaculatum oppositum
Agonum muelleri
Bembidion tetracolum
Stenolophus comma
Anisodactylus sanctaecrucis
Trechus quadristriatus
Harpalus pensylvanicus
Patrobus longicornis
Loricera pilicornis
Cicindela scutellaris lecontei
Pterostichus mutus
Bembidion rapidum
Agonum cupripenne
Bembidion obtusum
Geopinus incrassatus
Harpalus affinis
Chlaenius tricolor
Agonum placidum
Harpalus herbivagus
Syntomus americanus
Bembidion inaequale
Bembidion rupicola
Harpalus caliginosus
Harpalus puncticeps
Stenolophus lineola
Bembidion versicolor
Trechus discus
Agonum tenue
Bembidion americanum
Colliuris pensylvanica
Harpalus longicollis
Agonum cupreum
Anisodactylus rusticus
Bembidion nitidum
Bembidion salebratum
Chlaenius lithophilus
Omophron americanum
Harpalus faunus
Total number of beetles collected 5 3199
High tillage treatment HT2 from 19941997
Agonum muelleri
Clivina fossor
Loricera pilicornis
% Abundance
39.27
13.95
10.52
8.25
6.64
6.53
3.32
2.68
1.08
0.80
0.77
0.72
0.61
0.55
0.53
0.44
0.39
0.36
0.30
0.30
0.28
0.17
0.17
0.17
0.11
0.11
0.11
0.11
0.11
0.08
0.08
0.06
0.06
0.06
0.06
0.06
0.03
0.03
0.03
0.03
0.03
0.03
25.13
18.07
16.32
7.79
5.36
875
Appendix 1. (continued)
Species
% Abundance
Trechus discus
Bembidion obtusum
Tachys incurvus
Stenolophus comma
Bembidion rapidum
Agonum cupripenne
Harpalus herbivagus
Chlaenius tricolor
Harpalus puncticeps
Bembidion rupicola
Dyschirius politus
Agonum placidum
Bembidion transversale
Bembidion oberthueri
Agonum tenue
Amara rubrica
Bembidion nitidum
Tetragonoderus intersectus
Amara cupreolata
Anisodactylus nigerrimus
Bembidion transversale
Dyschirius globulosus
Agonum sordens
Amara aenea
Amara avida
Amara familiaris
Cicindela sexguttata
Harpalus affinis
Agonum darlingtoni
Agonum gratiosum
Agonum hypolithos
Amara littoralis
Dyschirius brevispinus
Dyschirius erythrocerus
Harpalus bicolor
Stenolophus lineola
Agonum cupreum
Amara carinata
Medium tillage treatment MT from 19941997
3.35
3.29
2.64
2.60
2.58
2.46
1.77
1.29
1.26
1.23
0.60
0.55
0.54
0.41
0.40
0.37
0.28
0.26
0.15
0.14
0.12
0.09
0.09
0.09
0.08
0.08
0.08
0.08
0.05
0.05
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.03
28.39
876
Species
% Abundance
Agonum muelleri
Clivina fossor
Stenolophus comma
Bembidion inaequale
Agonum placidum
Loricera pilicornis
Dyschirius politus
Agonum cupripenne
Amara avida
Agonum tenue
Bembidion nitidum
Omophron americanus
Harpalus herbivagus
Pterostichus mutus
Tachys incurvus
Chlaenius tricolor
Bembidion obtusum
Amara aenea
Bembidion rupicola
Harpalus affinis
Amara cupreolata
Dyschirius obtusa
Stenolphus conjunctus
Bembidion rapidum
Bembidion obscurellum
Stenolophus lineola
Agonum gratiosum
Agonum hypolithus
Cicindela formosa generosa
Calasoma calidum
Colliurus pensylvanica
Cicindela punctulata
Cicindela repanda
Dyschirius erythrocerus
Dyschirius brevispinus
Harpalus puncticeps
Harpalus erraticus
Harpalus bicolor
14.56
9.68
7.18
7.15
6.56
3.87
3.75
3.59
1.80
1.33
1.21
1.13
0.90
0.86
0.74
0.62
0.55
0.51
0.47
0.47
0.43
0.39
0.35
0.35
0.31
0.27
0.27
0.27
0.23
0.23
0.20
0.16
0.12
0.12
0.08
0.08
0.08
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
877
Species
% Abundance
Microlestes brevilobus
Stenolophus lecontei
Harpalus faunus
Agonum cupreum
Amara carinata
Bembidion sordidum
No tillage treatment NT from 19941997
Cicindela punctulata
Clivina fossor
Cicindela formosa generosa
Harpalus bicolor
Cicindela repanda
Harpalus herbivagus
Pterostichus luctuosus
Cicindela scutellaris lecontei
Stenolophus comma
Harpalus affinis
Amara aenea
Calosoma calidum
Harpalus puncticeps
Stenolophus conjunctus
Amara rubrica
Chlaenius tricolor
Amara avida
Agonum placidum
Harpalus erraticus
Notiobia picea
Amara cupreolata
Loricera pilicornis
Agonum cupripenne
Agonum muelleri
Amara latior
Calathus ingratus
Microlestes brevilobus
Pterostichus mutus
Agonum sordens
Amara convexa
Stenolophus lineola
Agonum tenue
Bembidion inaequale
Harpalus longicollis
0.04
0.04
0.04
0.04
0.04
0.04
21.95
17.51
10.28
7.08
5.74
5.31
4.81
4.55
3.92
3.71
1.96
1.62
1.08
0.95
0.89
0.84
0.70
0.65
0.65
0.65
0.58
0.50
0.50
0.48
0.34
0.28
0.21
0.19
0.17
0.17
0.15
0.10
0.10
0.09
0.09
0.09
0.07
0.07
0.07
0.05
0.05
0.05
0.05
878
Species
% Abundance
Omophron americanum
Patrobus longicornis
Tetragonoderus intersectus
Harpalus faunus
Amara familiaris
Amara littoralis
Bembidion nitidum
Bembidion rapidum
Geopinus incrassatus
Harpalus erythropus
Agonum hypolithos
Bembidion obtusum
Colliuris pensylvanica
Dyschirius obtusa
Notiobia terminata
Notiophilus aeneus
Stenolophus lecontei
Agonum piccolum
Amara carina
Bembidion sordidum
Harpalus funerius
0.05
0.05
0.05
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.03
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
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Biodiversity and Conservation 12: 851882, 2003.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

Assessing tillage disturbance on assemblages of

Received 14 June 2001; accepted in revised form 29 April 2002

Ferguson and McPherson 1985; Brust et al. 1986; Carcamo

diversity (Brust et al. 1986), whereas others (Carcamo

Materials and methods

Selection of agricultural fields

used. In T. aestivum, pre-emergence tank mixes of dicamba, MCPA, and mecoprop

Pitfall traps for the collection of carabid beetles

where H9 is the ShannonWiener index and S is the number of species.

where S is the number of species and N is the number of individuals.

where x is determined from S /N 5 [(12x) /x][2ln(12x)], S is the total number of

a x is the number of species predicted to have one individual, (a x ) / 2 those with

with x being estimated from:

where N is the total number of individuals.

To determine if our treatment data sets conformed to log normal distributions,

Species abundances were transformed using log 2 to generate intervals of octaves. S0

Table 4. Diversity indices for published carabid data.

Upland aspen forest

R is richness, A is abundance, H is the ShannonWiener index, E is the Shannon evenness index, BP is

Niemela et al. (1992)

Upland aspen forest

Halme and Niemela (1993)

House and All (1981)

Asteraki et al. (1992)

body size dependent (Muller

Brasil): I. Diversidade, abundancia

Fabreries Association des Entomologistes Amateurs du Quebec,

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