CARABID BEETLES AS INDICATORS OF TlUAOE DISTURBANCE

A Thesis

pnsented to

The Faculty of Graduate Studiw

of

The Univmity d Guelph

by

SVENJA BELAOUSSOFF

for the degrne of

Doctor of Philosophy

Much, ZOOO

@ Svenjr B.lroussM, ZOOO National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et Bibliographie Services senrices bibliographiques 395 Wehgton Street 395, nie Wellington OnawaON K1AW ûitawaON K1AONS canada Caneda

The author has granted a non- L'auteur a accordé une licence non exclusive licence aliowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seii reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur format électronique.

The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis wr substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Svenja Belaoussoff Advisor: University of Guelph, 2000 Peter G. Kevan

Canbid ôeetJes am ottln cons ide^ highly useful indicaton of distuturbance and envininmental stress. They am useful because they are taxonomically well known and are easiiy captured. Howcrver, there is no comprehensive study of carabids using a gradient of similar disturbances and a wide variety of ecological indices and models. I assessed the effect of tillage disturbance on the divenity within and btween carabid assemblages by using 14 diffemnt measums of dversity from samples collected on individual fams (i.e., treatments) in southern Ontark. The treatment8 npmsenteâ a gradient of Mage from no tillage to high tillage. I used 7 different divenity indices (ridiness, Shannon-Weiner, Berger-Parker, Q- statistic, Margalef, alpha and evenness). I examined divenity and abundance of beetle species and sizes by using deviation frorn divenity and abundance models. I used 3 different specirs divenity and abundance models (log normal, log series and geometric). For beetle sizes, I used 4 mctawms (maan Iength, divemity of Iengths, distribution of lengths, and distribution of species biovolume (length X width X depth)).

I used diffennt divenity indices to test the nuIl hypothesis that for al1 divenity indices there is no change in species diversity w'th increasing tillage disturbance. My hypotheses could not be rejected (psO.05) because no difference in divenity between fans was found.

These tests and resub mmamoborated by a meta-analysis of 45 published carabid data sets. Thus, rny own data. and those of othen, indicate that divemity indices are not useful for detecüng the effect of disturbance on unbids. Such a wide range of divenity indices king useà to assess the same data sets (original and published) had not been attefnpted before. Because of the negative findings obtained by using species divenity, I examined the novel hypotheses that diversity in beetle size, a functional measure of a beetle's place in an ecosystem, would be useful. I hypothesized that them would be decreasing divenity of beetle spedes sizes, increasing mean beetîe size and inmaring deviation away from log normal distributions of size with inudcising disturbance as npmented by the tillage practices on the 4 treatments. Because the nuIl hypotheses mmnot rejected, one anconclude that treatments, and the tillage practices, did not affect carabid community structure. The log nomial distributions of body size on al1 treatments suggest that carabid beetle assemblages fom a fundional group. Arguments of niche hieralchy and cornpetitive interactions can be invoked.

From the findings of the log nomality in sizes, I hypothesized that carabids are a taxonornic functional group by using distribution of species diversity and abundance. My results suggest that they are not because their divenity and abundance is not represented by log normal distnbutions. I found that carabid populations were geornetrically distributed in al1 treatments. Another hypothesis is that carabid populations would increasingly deviate from log norrnality with increasing tillage disturbance as repnsented by different treatments. This hypothesis could not be tested because the data never presented a mode in abundance. My results were compared to a meta-analysis of 45 published carabid data sets which confinns that carabids are not represented by log nomality even under undisturbed situations. Thus, carabids can not be used to assess the effects of disturbance by deviation from log nomality of specirs divemity and abundance.

To determine if lrvels of disturbance can be assessed by using other taxonornic groups I used a meta-analyses of non-carabid species assemblages to test for deviation from kg normal distributions of avenity and abundance. Funcüonal groups that should be targeted for distuhance studies aie keystone groups which include pollinatonr, and detritivores.

Underlying ecological principles to diversity and abundance would predict that some indicator groups may not show a change in divenity and abundance as the result of a distuhance. The disturbance under consideration may have been in place long enough for the group to be adapted to it, or the group may be alnady distuibed by an undetected force.

Although ecological indices and models are powetful tools that anbe used in assessing the effects of disturbance on ecological cornmunities, they must be used with care and in conjundion with the knowldge of the natural history of the seleded indicator groups.

Carabid beetle assemblages do not seern as useful as has been generally supposed because species w'thin the assemblage do not exploit resourcss in a similar fashion. Dedication

This dissertation is dadicated to my mothrr, Unula Belaoussoff. I would like to thmk my advisor, Dr. Peter Kevan, as ml1 as memben of my advisory cornmittee, rsprcially Dr. Vernon Thomas. Di. O. Bdan Allen and Ms. Jen Stone gave me valuable statistical advise, as did Dm. Steve Murphy, Medhat Nasr and Carlos Greco. David

Slater, Shamn Laws, Helen Sesward, Kym Welrtead, Tenl Feltz and Dan Guenther assisted mth my fieldwork. Also I would like to thank C. Schultz, M. Day, J. Gal, and the Perriman family for allowing my accort to thair fields. TABLE OF CONTENTS Abstrad Acknowledgements Table of Contents List of Tables List of Figures

Chapter 1: Introduction to Novel Approaches for Assessing the Effect of Disturbance on Diversity

Chapter 2: Assessing Tillage Disturbance on Carabid Assemblages with Diversity Indices

Abstract lntroduction Methods Seledion of agricultwal fields General experimental design Pit fall traps for the collection of carabid beetles Species identification Diversity statistics Statistical analysis Results General findings Carabid rictiness and abundance over transects Species diversity Meta-analysis of published carabid data sets Discussion General findings Species diversity

Chapter 3: The Effect O1 Tillage Disturbance On Body Sire Distributions Within Carabidae

Abstract Introduction Methods Results Discussion

Chapter 4: Carabid Beetles And Their Fit To Oiversity Abundance Models Under A Gradient Of Tillage Disturbance

Abstract Introduction Methods Results Discussion

Chapter 5: Fundional Groups Which Are Useful For Assessing Disturbance Via Deviation From Log Normal Distributions Of Oiversity And Abundance

Abstract Introduction Methods Results Discussion Chapter 6: Synthesis, Limitations And Future Directions For Assessing Disturbance

Synt hesis Limitations Future Direct ions

Appendix 1: Ecology of Carabid Beetles Species Collected From Southern Ontario Farm Fields 122

Appendix 2: Appendix 2: Key to Carabid Beetle Species Collected From Southern Ontario Fann Fields 152

Appendix 3: Data from Carabid Beetle Collections t 65

iii LIST OF FIGURES Fig. 1-1. Hypothetical rank abundance plots illustrating the typical shape of four species abundanœ models: geomeic, log series, log nomal and broken stick. 9

Fig. 2-1. Locations of the four ûeatmrnts (fams iepresenting different degrees of tillage) used in my study. 26

Fig. 2-2. Pooled abundance of species found in each treatment. 36

Fig. 2-3. Oendrograms of carabid assemblages based on Jaccard's index. 46

Fig. 3-1. Measures of a) depth vs. Iength and b) width vs. length for 10 beetles from 10 wnbid species. 60

Fig. 3-2. Canbid species length by sire class from pooled replicate data for treatments representing high tillage (HTI and HTZ), medium tillage (MT) and no Mage (NT). 62

Fig. 3-3. Canbid species biovolume by voîume class from pooled replicate data for treatments representing high tillage (Ml and HT2), medium tillage (MT) and no tillage (NT)* 62

Fig. 34. Mean carabid species length collected from treatments in each year representing high tillage (HTI and HTZ), medium tillage (MT) and no tillage (NT). 62

Fig. Cf. Rank abundance plots of carabid beetles wllected in my study on a logarithmic sale against the specirs' tank, in order hmthe most abundant to the least abundant species. 72

Fig. 4-2. Frequency distributions of carabid species cdlected from treatments npnsenting high tillage (HT1 and HT2). medium tillage (MT) and no tillage (NT), in relation to their abundanm. 72 Fig. 4-3. Rank abundance plots of carabid beetles from data sets induded in my meta- analysis on a logarithmic scals against the species' mnk, in order from the most abundant to the least abundant species. 73

Fig. 61. Rank abundance plots of data sets included in my meta-analysis on a logarithmic scale against the species' mnk. in order from the most abundant to the least abundant s~as. 85 LIST Of TABLES

Table 2-1. Vegetation which was adjacent to al1 replicotes (i.e., fields within farms) used in rny study. 24

T8bl0 2-2. Summary of phpical characteristics of the four tfeatments (fanns npresetnting different degrms of tillage) used in my study. 24

Table 2-3. Jaccard's index of similarity. 45

Table 2-4. Divenity indices for published carabid data. 50

Table 3-1. Width and depth rneasurements for 1O beetles of 1O different representative species. 60

Tabk 3-2. Mean carabid Iengths and divenity (Hl) of length colleded for replicate fidds from each treatment. 63

Table 4-1. Summary of Chi square analysis foi log series distributions for data sets in the meta-analysis of 45 data sets from 10 studies. R is richness and A is abundance. 76

Table 5-1. Diversity indices for species data in published studies. 84 CHAPTER 1t INTRODUCTION TO NOVEL APPROACHES FOR ASSESSING THE

EFFECT OF DISTURBANCE QN ONERSllY

Ecology is mted in the study of disturbance. It was because natunlists were concemed about the effects of industrial disturbances on British flora and fauna that the world's fint ecological society, the British Ecological Society, was fomed more than 85 yean ago (Sheail 1987). Although the impetus for the formation of the Society was snairopogenic distuhance, it has akays ncognized that natural disturbances also occur. Natuml distufbances wn the gamut from long terni changes over geological tirne to suddrn, catastrophic, events. Natural catastrophes anbe expected, even periodic, events (for example: fitmds (Steward & Popp 1987; Davy et el. 1990; Gaver 1997), or hurricanes (Zou et el. 1995, Vandenneer et el. 1998) and boreal forest fires (Wiser et al.

1997)) or unpredidable events (for example, meteoric impact (Glen 1990; Marshall &

Ward 1BW), or geomagnetic revenals (Crain 1971) and volcanic eniptions (Tsuyuzaki

1994)).

Expected natural catastrophes occur et somewhat regular intervals. Thus, there can be seledion pressures on biota, and species can adapt to this type of disturbance

(Scheiner & Teeri 1987; Davy et al, 1990). Not only can biota bewme adapted to expected ccitiistrophic distuibanœi, they can evolve to dapend on them for survival.

For example. Jacû pines nquim forest fins to npraduce (Lam& MacDonald 1998) and there are Collembola in the Central Amazon which need floods to complete their life cycles (Gaver 1997). In contrast, unexpected catastrophic disturbance events may cause extinction episodes because much of the biota is unadapted to those dramatic sudden disturbances. An example of this type of disturbance is the mass extinction of the majority of dinosaun Mich occuned Men a meteor struck the Earth at the end of the Cretaceous era (Glen 1990).

The types of disturbance ueated by human kings rnirror expected, and unexpected, natural disturbanœ events. Similar to natural disturbances, hurnan caused expected disturbances are generally les$ harmful to biota. Human actions mimic expected natural disturbances and thus the biota cm adapt to thern. Examples of biota which are adapted to human disturbance indude plants which are adapted to grassland buming (Morgan 1999; Peet 1999) and slash and bum agriculture (Roosevelt 1992;

Seibert 1993), plants and arthropods which anadapted to heavy metals (Morrey 1995 and Fraser 1980 respectively) and frogs which are tolerant to acid pollution (Pierce &

Sikand 1985). Human disturbances can also be equivalent to unexpeded, natural disturbance events if they are catastrophic. Unlike with expected disturbances, setection pressures resulting from unexpected disturbance do rot act on biota and thus, biota do not evolve to tolerate them. A few examples of catastrophic disturbances include the

Exxon Valdez oil spiH in Alaska (Davey et el. 1997), exotic species introductions such as opossum shrimp in Montana which caused the collapse of the salmon fishedes and subsrquent declines in Bald Eagle and Grizzly 808r populations (Hobbs & Huenneke

1992) and introduced Varna mites on bees (Mons & Flotturn 1997), and large scale destruction of habitats in North and South America (Clark 8 Enter 1984; Chadrasekaru 8

Flid 1998).

lncreasing intensity, frequency, and seventy of such catastrophic disturbances, ore of partkular concem to ecobgists busethey might Iead to mass ertincüon events of species thmughout the world (e.g., Ehrenfeld 1972; Meadows et al. 1972; Mitchell

1989; Wilson 1098). Species divenity is critical for the maintenance of ecosystern functions (se0 May 1975; Hutchinson 1978; Downing & Tilman 1994; Nilsson (L Grelsson

1995). The importance of species divenity for the functioning of ecosystems was proven by the serninal wo* of Tilman & Downing (1994) on grassland biodiversity. They found that study plots with greater plant species divenity were more resilient to the effects of drought than were plots with less divenity.

Ecologists often use changes in species diversity to detemine the eflects of disturbance because species divenity is an important component of any ecosystem

(May 1975; Hutchinson 1978; Magurran 1988; Hubbell pononal communication1).

However, it is necessary to use appropriate species assemblages as indicators of disturbance to detenine the effect of disturbance on species diversity. Species assemblages above the single species level are comrnunities, functional groups, guilds and taxocenes.

"Community" is used to describe any group of organisrns belonging to a number of different species that cwccur in the same area and interact through trophic and spatial relationships (Lincoln et a/. 1982). Ouild' is an assembiage of organisms that have overlapping roles in ecological processes in a particulor area (JaksiC 1981). For example, a seed disperser guild could indude ants, birds and mammals. "Functional

1 Penonal communication, Stephen Hubbell, Department of Ecology and Evolutionary Biology, Princeton University, December 1998. group' anôe definad as a grwp of not neassarily related species exploiting a common msoum base in a similar fashion. Within a functional group there is greater similarity in ecological msource requirements than within a guild, thereby implying that there is a (jreater degree of intespecific cornpetition (Colwell& Winkler 1984; Arthur

1984). Some examples of fundonal groups include pollinating bees (Kevan et al.

1997a1 Kevan 1999), livestock gut fauna (Iszak & Hunter 1WZ), and herbivorous macrokpidoptera (Lama et al. 1988; Hill et el. 1995). Unfortunately, the definition asuibed to guild by one author might be given to functional groups by another (see Root

1967; JaksiC 1981; Krebs 1985). To avoid confusion, in my dissertation, I use each term as it is defined above. Another terni to describe ecosystem components above the single species level h "taxocenen. This term is detined by Hutchinson (1978) as "an assemblage of specirs belonging to a particular classificetory unit or higher taxon, such as farnily, found in a given habitat. sharing resources". Taxocenes and functional groups are not necessarily mutually erclusive, and the species assemblages described by thern can potentially be described by more than one of the ternis. For example, a taxocene of pollinating bees can also be dassified as a functional group.

Although there is much similarity behwen the four types of species assemblages discussed above, aie most appropriate one to use for disturbance studies is 'fundional group'. 8ecause there is an overlap in resource requirements between species in a functional group, disturbances would affect those spedes by disrupting resources that they al1 use. Species within communities and guilds often depend on different resources, thus, s given asturbanœ would affect some of those resources but not all, and not alspecies would be equally affected. Because the species assemblages which make up communities and guilds are not egually affected by disturbances, they would not be as sensitive to disturbance as funcüonal groups would be. Species divenity and ibundanœ modds (ses below) provide one method of measuring disturbance but should be applieci only to funcüonal groups.

Fundional groups evolve to occupy the available niches within ecosystems.

Ecosystems comprise producen, consumen and decomposen which are linked together by energy, producüvity and nutrient cyding (Krebs 1985). Lakes, estuaries, marshes, forssts, and tundra al1 anbe wnsidered to be different types of ecosystems.

Ecosystems are not only natuml, undisturbed amas, they can also be arüficially dominated by human activities. Agricultural fields, for example, cmbe considered as ecosystems because they have the properties of an ecosystem (e.g., producen and consumen) (Kevan et al. 1997b). When an ecosystem, whether it is natural or dominated by human activities, is disrupted, the îunctional groups within it are affected.

Although them ana number of approachrs to assesring the effects of disturbance on spedes, induding flagship species, indicator species, keystone species, and resilience, I have chosen to focus on two different approaches to assess the effect of disturbance on species within fundional groups: divenity indices, and deviation from divenity and abundanœ models (e.g., species sire diversity (see Chapter 3) and spedes divcmity and abundance (see Chapter 4)). The prirnary reason I chose to use those approaches is because they are useful for addressing questions regarding fwictional gmups (see below). Of those two approaches, divenity indices are more ftequently usdby ecologists. Divenity indices are generally simple to calculate (with some exceptions, ag., the Q-statistic; also see Magurran 1988 and Chapter 2), especially in cornpanson to the mathematics and statistics behind diversity and abundanco rnodetr (ree below and Chapter 4). Although there are many different diversity indices, they fall into one of two cetegories: species richness indices, and indices basad on the proportional abundances of species (see Magunan 1988).

Richness indices are a measure of the number of species, or individuats, in a defined sampling unit (e.g., nurnber of species, density, abundance). Indices which measure the proporüonal abundance of species combine species richness and evenness (the component of species divenity that measures the relative abundance of species) into a single figure (ree Magurran 1988) (e.g., Shannon-Weiner, Berger-Parker, Q-statistic,

Margalef, alpha and evenneu itself). These indices tend to be biased either towards species richness (e.g., alpha, Q-statistk) or dominance (e.g., Berger-Parker) (Magunan

1988).

Divemity indices cm k useful because they provide rapid, and easily calculated, ecological measums. Also, becsuse many researchers use indices, it is possible to make some cornparisons between similar studies which use the sarne indices.

Neverthelers, although they are wmmonly used, there are three main shortwmings of indices. Fint, they can be used only for cornparisons between sites, or on sites over time, or boal (for example, studies on the affect of micring waste on fish (Comell et al.

1978), landfill mdamation on invertebrate populations (Judd & Mason 1995) and altitude on spider populations (Otto & Svensson (1982)). Second, divenity indices an, only statistical artifices and do not have any intrinsic biological meaning (see Southwod 1978;

Magurran 1988). Third, the use of different indices Mai the same data con result in diffemnt condusions (see Chapten 2 and 5).

One way to overcome the shortcomings of divenity indices is to have a diversity measure which is 'stand-alonen and does not need inter-site or inter-üme comparisons.

Wth such a stand-alono mwswo a theoretical standard against which to make comparisons is nquired. Such a standard should k, based on community structure, and ewlogical and evolutionary theory, to circumvent the shortcoming of being a statistical artifice. By using a standard, it would be possible to compare the structure of a funclional group against the standard and essess whether or not the fundional 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 (also see below, and Chapten 4 and 5). Divenity and abundance models, and distributions of sire within functional groups, provide a good starüng point to find the standards needed for stand-alone measures of diversity because they anbased in ecdogical and evolutionary theory.

Although diversity and abundance models have been present in the ecological literature for over 80 yean (e.g., Raunkiw 1918). divenity indices are more frequently used by ecologists to examine the effed of disturbance on species divenity (Southwood

1978; Magurran 1988). Relationships describing the diversity and relative abundance of spdes in a community wem origindly suggested by Raunkiw (191 8), and Vien again by Motomura (1932). Although diennt divenity and abundance models have been developed (e.g., Mandelbmt's (1977) Zipf-Mandelbmt model; Dewdney's (1997) logistic

J and Hubble's (personal communication) zemwm multinomial), four principal distributions of diversity and abundance (MacArthur's broken stick rnodel, geometric series, log series and log normal) rernain central to discussions about diversity and abundance rnodels (Magumn 1988). Beceuse those four distributions are commonly considered, they are the ones used in my dissertation. Furlhemore, sorne of the altemate models am not based on ewlogical principles (e.g., the Zipf Mandelbrot model is based on information theory (Magumn 1988)), or are rmpirical fits of data which cannot be separated from log nomal distributions (e.g., the logistic J model

(Dewdney 1997)). When plotted on a rank abundance graph (Figure 1-1) the four models represent a progression ranging from the geometric series in which few species are dominant with the remainder uncornmon, through to the log series and log normal distributions with species of intemediate abundance becoming more common and ending in the conditions reprssented by the broken stick mode1 in which al1 species are approximately equally abundant (Magunan 1988).

The broken stick model reflects an equally divided niche spaœ (MacArthur 1957).

Its biological meaning is not obvious (Hutchinson 1978; Magurran 1988) although, rmpirically it is most likely to occut in groups of somewhat mobile animals living in a homogeneous environment Unlike the other models, this model reflects an equitable state (Magurrsn 1988) and there is no cornpetition between members of the fundional group. The value of the Sroken stick model has been challenged (Wilson 1993), and there are few exarnples of where it occun (e.g., passerine birds; MacArthur 1960; minnows; King 1964).

Both the geometiic and log series distributions ara believed to result when niche Fig. 1-1. Hypothetical rank abundance plots illustrating the typical shape of four species abundance models: geometric. log series, log normal and broken stick. In these graphs. the abundance of each species is plotted on a logarithmic scale against the species' rank in order from the most abundant to the least abundant species. Taken from Magurran (1 988). pmemption occun (Le., a few species pm-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 th8t geometric distributions am found in species-pwr (often hanh) environments, in the very rady stages of succassion (Whittaker 1985, 1970,1977) or under disturbed conditions

(e.& Kempton & Taylor 1974). As succession proœeds, or as conditions improve, species abundance distributions becorne log series (Magurran 1988).

The log nomal mlationship describing air divenity and relative abundance of species in a community is one of the oldest proposed (Raunkiaar 1918; Motomura 1932). ln comrnunities described by this distribution there am few common spedes, more less common species, and few rare species. It was Preston (1948) who firot tested the idea that species divenity and abundance can be described by log nomally distributed frequencies. Since then aie model has been shown to fit many biological communities

(e.g., diatoms (Patrick 1988); soi1 arthropods (Hainton & Byen 1954); macrolepidopten (Kempton & Taylor 1974; Laroca et al. 1989; Hill et el. 1995), birds and mammals (Preston 1962); bees (MacKay & Knerer 1979; Tepedino & Stanton 1981;

Stubblefield et al. 1993; Kevan el al. 1997a) and Salmonelle strains in livestock (Izsak &

Hunter 1992)).

The log normal model of divenity and abundance is more than a statistical propetîy of sampling (d. May 1975). Sugihara (1980) suggested the niche space of a taxonomically rebted gmup is sequentially split by the constituent species. and the fraction of the niche space apporüoned to a species is proportional to its abundance. The log normal model, as described by Sugihara, is also known as 'Sugihara's sequential breakage modeln. Until Sugihara (1980) broached its biological rationale, the log normal mode1 was deemrd rnerely to be a statistiwl consequence of the Central

Limit Theorem (MacArthur 1980; May 1975; Tokeshi 1993). The model ascribes an organizational pattern for a fundional group, and is based on the following considerations (Sugihara 1980):

1. Vie existence of an hieratchical niche structure in the functional group;

2. the underlying stnicture of niches is reflected in the pattern of abundance;

3. it is applicable to functional groups.

A hietaichical niche stnictun implies that each species making up a functional group is different from the othen and none is redundant. Tilman 8 Downing's 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 define quantitatively a functional group, and 2) the need for en indicator of disturbance which is free from temporal, or spatial, cornparisons of species. Generally, reseanhen classify species assemblages as fundonal groups, guilds or tanocenes because the species within the assemblage are dosely related, occur together in a particular habitat or seem to have similar ecdogical niches (sw above). Cornpetition occun between constituent species of functional groups. Thus, it is possible to assess them quantitatively. It is useful to use quantitative methods to describe a functional gmup in order to give the definition greater scientific validity thon if it is merely based on subjectively grouping species togethet (JaksiC l98i). By using fit to the log normal mode1 under undistuibed conditions it may be possible to assess quantitatively if a functional group is present. However, if ais way of describing a functional group is not used with caution, it is possible to place more than one fundional group together and still obtain a log normal distribution of divenity and abundance (see Chapten 4 and 5). Even so, this way of describing a luncüonal group is an initial, and much needed, quantitative approach to determining if a colledion of species form a functional group. Researchen using log nonnality to define functional groups have to be conssrvative, and cognisant of the ecological backdrop in which log nomality is used.

Preston (1980) proposed that the degree of deviation of diversity and abundance from log nomslity can be used as an index of the degree of disturbance. This is the second approach to measunng the effect of distuturbance on functional groups (see above). The idea that deviation from log nonnality might reflect the effect of disturbance has been tested with diverse assemblages of marine organisms (Gray & Mina 1979;

Gray 1979. W83). Those studies supported the hypothesis that then, would be deviation frorn log nonnality, even though the organisms that were sampled did not bekng to a single fundional group. Thousands of individuals were collected, and it is likely that several functional groups were each well represented and were al1 similady affected by the pollution exarnined in Gray (1983) and Gray & Mina (1979). To date, studies which have associated departure from log nomal distributions of divenity and abundanw with disturbance have been on pollinating bees (Kevan d al. 1997a),

Lepidopten (Lama et a/.1989; Hill d el. WQS), gut bacteria in Iivestock (Izsak &

Hunter 1W2), and soi1 arthmpods (Higvar 1994). Using drviation from species divenity and abundance relationships to detemine whether or not a biologieal community is disturbed is mtmversial (Lambshead et al. 1983; Shaw et el. 1903; Nelson 1987). In general, the major critics of the method (e.g., Nelson 1987) have lumped unrelated, andlor only partial, fundional gmups together Viemby dismissing the approach without understanding why it did not work.

Another approach to measuring the effect of disturbance on a functional gmup is deviation from log normal distributions of size. This appmach is related to log normal distributions of diversity and abundance (Brown 6 Maure? 1986; Siemann et a/. 1998; see Chapter 3). Cornpetitive interactions between species leads to log normal distributions of species divenity and abundance. Likewise, cornpetition and the subsequent division of energy and resources within a functional group in undisturbed habitats con lead to log normal size distributions of species within habitats (Harvey 8

Godhgy 1997). Within a functional group there would be a few large species able to gather many of the resources (Morse 1980), a few smallet species utilizing resources that are energetically inefficient for larger species to collect (Pearson 8 Mury 1979) and a large number of intermdiately sized species. When, and where, there is a disturbance which interferes with resource availability, the cornpetitive interactions between species in a functional gmup is disrupted, and a shift away from log normal distributions of size would occur. This approach to using rnodels and size distributions is not reported in the literature.

Funcüonal groups exposed to unexpected disturbance events are expected to deviate fmm log nomality. Howevrr, such deviation merely raises a red flag which indicates that the funcüonal gmup in question may have been disrupted (Kevan et el. 1997a; Belaoussoff & Kevan 1998). Atcording to some reseanhen, log nonnality is a characteristic of cornmunitirs in equilibrium (Stenseth 1979; Ugland 6. Gray 1982).

Howcwer, a study on birds show that undistuibed fomt habitats are not necessarily at equilibrium (Gee & Giller 1986). Thur, situations exist in which functional groups in undisturbed habitats might deviate from expected log nonnality. 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 fundional groups is a novel approach. More work is needed to sort out the camplexities of functional groups and deviation from log nomality as an indicator of disturbance (Hill & Hamer 1998).

My study examines methods to define quantitatively functional groups with respect to Carabidae (i.e., carabid or ground beetles) on fanns representing a gradient of Mage disturbance (see Chapter 2). Carabidae need to be examined as a potential group for assessing disturbance because they are considered to be sensitive indicaton of environmental change (e.g., Dritschillo & Wanner 1980; SekluliC et al. 1987; Eyre &

Rushton 1989), they are abundant within a wide range of ecosystems (see Thiele 1977), they are relatively eesy to catch in large numberrr (Thiele 1977), the taxonomy of many species is well known (Lindmth 1961-1969) and expert identification is accessible.

Furthemon, more is known about the ecology of carabids than of almost al1 other insect families (Thiele 1977; Lamchelle 199Q also see Appendix 1).

It is poorly undentood how carabids are affected by agricultural disturbance. Pmvious studies which have attempted to assess conseqwnœs of tillage on carabid diversity wen not condusive. For example, several studies have show carabid divenity is gmater in fields under minimum üllage than conventional tillage (DritschiHo & Wanner

1980; House & All1981; House (L Stinner 1983; Feguson & McPhenon 1985; Bnist et

81. 1986; Cdrcamo 1995; Weiss et a/. 1990). However, other studies either indicate tillage does not affect canbids (Tyler & ENis 1979; Hokkanen & Holopainen 1986; Dritschillo & Erwin 1982; Kromp 1989; Mack & Budvnan 1990; Tonhasca 1993) or even that carabids are more diverse in conventionelly tilled fields (Edwerds 1976; Bamey & Pass 1986a). Some researchen also have found rotation and crop type affected canbid divenity (Lovei 1984; Bnist et al. 1986) whereas othen (Charno et al. 1995) did not. One problem with most of these studies is that the only divenity measures that were used were richness and abundance. Although these measures have intuitive appeal, they do not provide information about changes in the relative proporüons of species, i.e., community structure (Magurran 7988). To find out how useful carabid beetles are as indicator species, it is necessary to explore fully their potential with different measures of divenity (see above). Although diversity indices have been used with canbid beetles on other studies (see refenmces in Chapter 2), the beetles need to be examined in the context of deviation from log nomal distributions of divenity and abundance, and size.

In my dissertation I examined the following:

1) In Chapter 2, 1 used 7 diffennt divenity indices (richness, Shannon-Weiner, Berger-

Parker, Q-statistic, Margalef, alpha and evennsss) to address the nuIl hypotheses that

for al1 divenity indices then is no change in divemity with inmaring tillage disturbance

as npresented by different fams (ses Chapter 2), and by different crops, within famis

or years. I tested îhese hypotheses with repeated measures analyses. These tests are supplemented by a meta-analpis of 45 published carabid data sets from 10 studies that represent carabid communities collectecl fmm undisturbed to highly disturbed sites. A novel aspect to this chapter is that a wide range of divenity indices are used to assess the same data sets.

2) In chapten 3 and 4, 1 examined the hypothesis that canbid beetles are a functional group, and whether the group, functional or not, is sensitive to disturbance. These two chaptes represent two different approaches to the hypothesis, with Chapter 3 using distribution of divenity of beetle size and Chapter 4 using distribution of species diversity and abundance. Quantitative definitions of funcüonal groups are needed, yet have not kenexplored in the published literature.

3) In chapter 3, 1 examined the novel nuIl hypothesis that there is no change in size distribution of carabid beetles with increasing tillage disturbance as represented by different fanns (see Chapter 2). 1 examined size distributions with 2 different methods: biovolume and mean Iength. I also used mean lengths of carabids on each fam, as well as a novel modification of the Shannon-Wiener index which used length. With the modification of the Shannon-Wiener index spa*estIengths were considered rather than taronomic classifications of species.

4) In chapter 4, 1 examined the hypothesis that canbid assemblages would increasingly deviate fmm log normal distributions of divenity and abundance wlth increasing tillage distuturbance as represented by diffemnt farms (sm Chapter 2). These tests are supplemented by a meta-anaiysis of 45 published canbid data sets from 10 studies that represent csnbid communities colleded from undisturbed to highly disturbed sites. This approach has not been taken bafore with carabid beetles. 5) In chapter 5, I examined nonarabid assemblages by using published data sets from a range of organisms (induding spiden, Lepidoptera, tundra mites, bees and diving beetles) under different disturbances (ranging fmm clear cutthg, industrial pollution and agncultural activities) to furthet examine the hypothesis that fundional groups deviate from a log normal distribution of divenity and abundance when exposed to disturbance. CHAPTER 2: ASSESSING TILLAGE DISTURBANCE ON CARABID ASSEMBLAGES

MTH DIVERSIlY INDICES

ABSTRACT

Many ecdogical studies have used divenity indices to assess disturbance.

However, most have used only one or two indices. Exceptional papen have used three indices, but none has used multiple indices. I used seven different divenity indices

(richnesr, Shannon-Weinri, Berger-Parker, Q-statistic, Margalef, alpha and evenness) to test the nuIl hypothesis that for all divenity indices there is no change in divenity with inueasing tillage disturbance as represented by different fams (treatments), and that thete is no difference in diversity with different crops or years. I tested these hypotheses with repeated measures analyses. I was not able to reject the nuIl hypothesis that there is any divenity difference among fanns. I also found that no one divenity index was better than any other a detecting disturbance. These results were supplemented by a meta-analysis of forty five published carabM data sets. The meta-analysis supports the conclusions that divenity indices are not useful for detecthg the possible effect of disturbance on carabid beetfes.

INTRODUCTION

This chapter is part of theapproaches to addressing whether or not fundional gmups (see ddnition in Chapter 1) are useful as indiatom of distuturbance (see

Chapters 3 and 4 for the oaier appmaches, also Chapter 1). Of the three appmches, divenity indices are most wmmonly used by ecologists. Many ecological studies have used divenity indices to assess disturbana. Howevet, most have used only one or two indices, typically fichnesr Mai the Shannon-Wiener index and less wmmonly.

Margalds index (e.g., Comell et al. 1976; Niemela et al. 1992; Judd & Mason 1995;

Kithara & Fujii 1994). Exceptional papers have used aime indices (e.g., both Doane &

Dondele (1979) and Hill et al. (1995) who used three), but none has used multiple indices that ernbrace concepts of dominance, richness and ecological information (see

Chapter 1). I have used seven common indices (richness, Shannon-Wiener, Margalef, alpha, Q-statistic, Berger-Parker, and evenness) to evaluate the same data base.

Studies which use multiple indices are needed because it is not known which ones pmvide the most sensitive indicator of disturbance. This approach has, so far, been overlooked,

ln my study, I examined the effect of Mage disturbance on carabid beetles.

Tillage, aie procesi of tuming the soi1 and incorporating organic litter into it, is an important aglicultural pracücs. Shifts fiom high tillage to no tillage practices have been occurring throughout North America since the 1974s (House & Stinner 1983). The main types of tillage used in southem Ontario are: high (mold board plow), medium (chisel plow) and no (wnservation) tillage. In Ontario, no tillage is aie hast cornmon of the thme practiœs (sinœ the mid 19Ws il has been used on only 10-1 5% of fams

(Clarence Swanton, personal communication2). High Mage penetrates approximately

2 Persona1 communication, Clarence Swanton, Department of Plant Agriculture, University of Guelph. Novrmbw 1999. twice as fer as medium tillage. No tillage involves seeding with minimal disturbance

(Oryokot et el. 1997). Fields under high and medium tillage, in contrast to those under no tillage, lad< distinguishable soi1 horitons (AlIrnarus & Dowdy 1985), have less organic detritus on top (AlIrnarus & hvdy 1985), are much drier (House & Stinner 1983;

Allmanir & Dowdy 1985), and exhibit more soi1 compaction (House & Stinner 1983;

Tonhasca & Stinner 1991). Because increasing Mage decreases the number of micro habitats available to the beetles and decreases the abundance of their prey (Stinner et el. 1988; House & Parmelee 1985), carabid divenity is likely to be effected by tillage.

Although there are negative consequences of tillage, Neave (1992) showed that the fauna of tilled fields rewver from tillage disturbance more quickly than the fauna of old fields.

I used carabid beetles which were collected from four farms that represent a gradient of tillage disturbance, and I supplemented my findings with a meta-analysis of forty five published data sets. My nuY hypotheses for ail divenity indices are that then would be no change in divenity with increasing tilkige, or other disturbance. In addition to the seven different diversity indices (richness, Shannon-Weiner, Berger-Parker, Q- statistic, Margalef, alpha and evenness) I used Jaccard's index to examine the relative similanty of the species composition among fields, fans and yean. The other indices do not compare the achial species presence and absence, but pertain to divenity per se. I also examined the spetcies richness and abundance over time (season) and by trap position. Additionally, I sepantd out the native and exotic species because colonizing rxotic spedes often ocair in higher abundance than native species in disturbed rnvironments (Kmbs 1985). METHODS

Selection of Anricultural Fields

My study was part of a Tri-Council agroecosystern research project which was based at the University of Guelph. To be part of that project, a constraint of my wodc was that it had to be located within the Grand River Watershed in southem Ontario. I selected my sites in conjunction with On. S. Murphy and B. Forbes, with whom I worked closely because we al1 designed our research projeds to be complementary. Whereas my study was to examine carabid beetles in a gradient of tiHage, Dr. Murphy's study was to examine weed plant divenity and Or. Forbes' study was to examine fion in riparian areas which anadjacent to agricultural fields. My experimental design was decided upon after intensive consultation with Dm. Murphy and Forbes. Ors. C. Swanton, B.

Smit and D. Rapport, with whom Drs. Murphy and Forbes were working, my advisory cornmittee , as well as Dr. C. Greco (a post doctoral feliow with Dr. Kevan), were aiso consulted about my field research design. For our research, we could not use experimental fanns because we would have had plots smaller than farm fields. Further, we would have had to wait for yean befon kingable to begin the expenment because the environmental effeds of a new tillage pnctiw take about 3-10 years to establish

(House & Stinner 1983). Also, because Carabidae are mobile, small plots in experimental farms would not have been usefut for addressing questions about the effect of tillage on their divenity. On an experimental fann it would not have been possible to have 8 number of fields with borden that had similar vegetational composition, or other lldswith the same tillage prstace. Although it might be ideal for the sort of research we undertook to have had several large fams, each with a divenity of tillage practices, that is not pracücal. It is also not feasible to as& CO-operatonto change their fanning practices to meet the needs of experiments like ours. Even if a CO- operator agreed to change tillage pnctices, there still wwld be a waiting period of about

3-10 years for the environmental effects of the new tillage regime to establish.

However, the major problem with the experimental design which was used in my study is that there is no true replication, rather the fields within farms are pseudoreplicates. This problem is a flaw within the design which I used and any one designing similar experiments should be Pware of the flaw. Nevertheless, the justification for using that design is ouUined below.

We began with over 100 willing CO-operatonwithin the designated area, and visited over 30 of them. To ensure that any effects of tillage which we might find would refied the current tillage regime, we required fanns on which the same tillage practice had been in use for a minimum of five yean. This constraint was most limiting for the seledon of a no Mage fann because only 10-1596 of fannen in the Grand River

Watershed had bwn using no Mage for at least 5 yean before my study cornmenced

(Clarence Swanton, penonal communication3). We wanted to control for as many variables as possible and originally ended up with 3 fams which were highly comparable except for tillage practice (see below). In the second year of the study we found an additional high tillage fann which neighboured the no tidage fam. As a group, we

3 Penonal communication, Clarence Swanton. Department of Plant Agriculture, Novernber, 1990. agreed that al1 the fields to be used were to be bordered on 1 side by a wood lot to assure conformity of edge vegetation on at least one side of the field. On. Murphy and

Forbes surveyed the vegetation in the wood lot and field margins surrounding al1 potential fields, and we agreed only fields sunounded by similar vegetation could be used to assure comparative rigour (Table 2-1). All of the fields that were used were also nstricted by being more or Iess the same redangular shape (approximately twice as long as wide) and area (approximately ten hectares). The slopes, aspects and soi1 types of al1 fields also had to be similar. However, one of the high Mage fams had slightly more loamy soi1 than aie oaier fams (Table 2-2). Soils were tested as to type using methods described by Spurr & Bames (1980). All fams had to have similar applications of the same pesticides and fertilizen (Murphy & Swanton 1999). The average corn heat units for each of the farms used ranged between 2650 and 2750 (Brown 8 Bootsma

1993).

In summary, the following 12 wnstraints had to be met in fam and field choice:

1. The fans had to be within the Grand River Watershed.

2. The fams had to use one of 3 tillage practices: high, medium or no till.

3. The fanns had to be useful for the research of On. Murphy, Forbes as well as for my work.

4. Crop rotations on the fams had to be simibr.

5. Agrochemicals which were used on the fams had to be similar.

8. Slopes and aspects of the farm fields which were used had to be similar.

7. Soil types of aie fam fields which mmused had to be sirnilar.

8. Vegetation and fields sunounding the fam lldswhich were used had to be similar. Table 24. Vegetation which was adjacent to al1 replicates (i.e., fields within farms) used in my study.

MOWED MARGINS WOOD EOGE WOODS (MAPCE-BEECH) Duacus carota Poa prafensis Acer saccharum Canyza canadensis Phleum pratense Carpinus caroliniana Amaranthus retroflexus Dacty/is glomerata Ostrya virginiana Ambrosia artemisiilola Bfornus inermis Viburnum acerfolia Elytrigia repens Equisetum spp. Alliaria petiolata Eryîhoniurn americanum Claytonia virginiana Geraniurn robertianum

Table 2-2. Summary of physical characteristics of the four treatments (farms representing different degrees of tillage) used in my study. HT1 represents the mold board Mage (i.e., high thage) treatment in Oxford County and HT2 represents the rnold board tillage treatment in Wellington County. MT is the chisel plowed (Le., medium tillage) treatment in Waterloo County. NT is the no tillage treatment in Oxford County. Slope is in degrees. Soil type was deterrnined using methods of Spurr and Eames (1980). Three crop rotation is corn-soy bean -wheat, two crop rotation is corn-soy bean. Replicate refer to field within farms.

Tillage type Location Treatment Replicate Soil type Slope Rotation No till Oxford NT NT-1 sandy-loam 0.25 3 crop NT-2 sandy-loam 1.50 3 crop NT93 sandy-loam 1.25 2 crop NT4 sandy-loam 1.O0 3 crop

NT-5 sandy-loam 1.25 2 crop t Moldboard Oxford HTI HTS-1 sandy-loam 1.25 3 crop HTI-2 sandy-loam 0.25 3 crop HT?-3 sandy-loarn 1.25 3 crop Chisel Waterloo MT MT-1 sandy-loarn 1.50 3 crop MT-2 sandy-loarn 0.75 3 crop MT-3 sandy-loam 1.50 3 crop MT4 sandy-loam 0.50 3 crop MT95 sandy-loam 0.50 3 crop MT4 sandy-loarn 0.25 3 crop Moldboard Wellington HT2 HT2-1 loam 0.00 2 crop HT2-2 loam 0 .O0 2 crop HT2-3 loam 1.75 2 crop HT2-4 loam 1.75 2 crop

24 9. F#d shape and size (about 10 ha) had to be sirnilar.

10. Coipontors had to be Wling to let us work on their fami fields for the dumtion of ouf studies.

11. Faming pracücb for each fam used had to br constant for 5 yean prior to our research.

12. Farming pracüce for each fann used would nmain constant for the dumtion of our studies.

The fams Hihich ultimately wen seleded differed primarily in degree of tillage which the farmer used. We chose the fams which, based on past pmctices, most resernbled one another, and we are confident that the main difference between them is in tillage. All memben of my advisory cornmittee concurred that the fields were highly similar with the exception of tillage.

With the above sedes of stringent requinments, we ended up with 4 farms (with a total of 18 fields representing 3 different fanning systems) in Wellington, Waterloo and

Oxford Counties, al1 withh 30 km of one another (Figure 2-1). With such lirnited choice, we decided that use of repliute fields within the similar fams would be our experimental design. The sampling mirne had to k kept to r pncücal levd to be manageable by few people. The km$used in my study ancomrentionally (mold board) tilled, chisel plowed or no till. For the rernainder of my dissertation, fields within fams which were tilled with a mold board plow are called HT (i.e., high tillage), the chisel plowed fields are called MT (i.e., meâium tilkge) and the no tillage Mds are called NT (i.e., no tillage). In

Oxford county, HT1 is adjacent to NT. ûther variables am aop typ (*ter wheat, soy Fig. 2-1. Locations of the four farms which represent different degrees of Mage in southern Ontario. HT1 and NT were in Drumbo, Oxford County, HT2 was in Elora. Wellington County, and MT is in Ayr, Waterloo County. See text for descriptions of farms.

Elora Lake Ontario .a Av Drumbo bean and corn) and aop rotation (corn-soybean, corn-soybean-wkiter wheat). The HT farrn in Oxford County (HT1) is located on Bedford-Blenheim Road. Drumbo. The NT fann in Oxford County (NT) is located on County Road 22, Drumbo. The MT fann in

Wateiloo County (MT) is located on County Road 97, Ayr. fans had the constraining variables we used tor the NT and MT fams. In 1995, an additional high Ullage coopemtor was included (in Oxford) (HTI) whose fam had 3 fields which were within the constraints outlimd above, and the HT2 cooperator gave me access to mon, fields, of which only 2 additional fidds which were within the constraints outlined above,

Genenl Exmrirnental Desian Ialso see below under Statistical Desian)

In my study, Icornpared carabid diversity between fams without replication of fann types, but rather with replications of fields within fam. Then is a precedence in the published ewlogical literature to conduct studies which are comparative but without replication (e.g., Kevan et al. (1975) on bees; Comell et al. (1976) on fish; Otto &

Svensson (1982) on spiders; Niemela el al. (1992) on carabid beetles; Witrnan (1992) on coral reefs; Clark et al. (1994) on carabid beetles; Kitahan & Fujii (1994) on

Lepidoptera; Charno (7995) on carabid beetles; Hill et el. (1995) on Lepidoptera; Judd et al. (1995) on invertebrates; Paoletti et al. (1995) on soi1 macroinvertebrates; and

Petenen (1995) on Collembda).

Although I concedr that an exprimenW design that would have many replicates of each fann type would be ideal, the design Dm. Murphy and Forbes and I used for ou? studies are valid and appropriate. We chose to use fields within fanns as replicates, and famn which mpresent different degrees of tillage as our treatments. To test our hypotheses that divenity would be affected by tillage, we wmpared divenity of our indicator groups among treatments. Appropriate, and robust, stotistical analyses for our experimental design were used (see below under statistical design).

Two type of experiments are wnducted within the disciplines of ecology and evolution (Scheiner 1993). The fint are manipulative experiments which include laboratory studies of behaviour, manipulations of growth conditions and imposition of experimental treatments on naturel environments. The second are comparative, observational studies. These are especially important for ecologists, and evolutionary biologists, because it is otten not possible to manipulate conditions in order to address hypotheses (Scheiner 8 Gurevitch 1993).

Although observational field experiments are necessary tools for ecologists and evolutionary biologists, as with every experimental design there are difficulties assodated with them. Those experiments are expensive, tnie replication may be unattainable (e.g., Carpenter IWO), there is 'noise " in the data because the environment is heterogeneous at many scales, and field measurements are crude cornpared to those obtainable in the laboratory. However, when conditions between sites are similar, and them is only one major factor which is different between sites, it is possible to make meaningful cornparisons between them. One of the aiteria ecologists use when looking for field sites is that they are as similar as possible except for the parameter that is to be tested. An integml part of training for al1 dogists is learning to select tield sites.

Pit Fall Tra~sfor the Collection of Carabid Beetîes

Carabids were collected from 1994 to 1997. Each nplicate (i.e., field) had a transect of 7 single pit fall traps. The traps were 8 cm deep and 15 cm in diameter white plastic tubs. The location of the transed 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 metres fmn the 3 other edges of the field. For each transect single traps were placed at the aop edge border, 1, 2,4, 8, 10 and 32 metres into the crop.

Additionally, in 1996 and 1997,2 bamer traps were put into place into soy and corn replicates to inwease sample sites. The fint trap was placed 1 metre from the crop edge nearest the woodlot end the second trap was placed 32 rnetres 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 bamer traps were fint tried out in 1996 ki NT and HT1 fields, and then put into al1 non-wheat replicates in 1997.

The 2 bamer ûaps were also placed aibitrarily along a transect in the replicates, but wen, separated from the single tmp transech by at lead 30 meters. Bamer traps consisted of 2 m long X 15 cm high woodrn boards which wem 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 wuld be placed flush aoainst the board when the trap was put into the soil. Two and a haIf meter high bamboo poles were placed beside each trap to indicate its location. Each year, the traps were installed in June, after corn and soy crops were planted and weather pennitted. The traps were filled with water and two drops of detergent to break surlace tension. Once a week al1 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 (1981-1 969) was developed to help with the identification of the species found in the collection (Appendix 2). Identifications were verified against specimens in the carabid collection in the University of Guelph insect museum and confimied with Andrew Applejohn (University of Guelph) and Henri Goulet

(Biosystematic Research Centre, Ottawa). Voucher specimens have been deposited in

Vie Univenity of Guelph insect museum. Appendix 1 provides a literature summary of the ecological characteristics for the carabids collected in my study. Histograms were prepared to illustrate the abundance of carabid species in each treatment.

My experiments wem not designed to identify a single indicator species, or even a wmplex of species. Rather these experiments were designed to look et the overall patterns of change refiected by disturbance on carabid beetle assemblages. Rie diversity indices which wem calculated indude dchness, the Shannon-

Weiner index, evenness, the Berger-Parker index , me alpha index, Q-statistic, and the

Margald index (Magurran 1988; Southwood 1978; Taylor et el. 1976). These indices were calculated for carabid beetle assemblages collected from replicates in HTI, HT2,

MT and NT. Jaccard's similarity index was also calarlated to determine within year similarities between and within replicates in the treatments.

Shannon-Wiener Index ( H*)was calculated as:

H'= -P( p,lnp, ) Ww where p,=N/N. N, is the number of individuals of Vie ith species in the sample and N is the total number of individuals,

Shannon evenness was then calculated using the formula:

E= H '/lnS @W.(2) where H* is the Shannon-Weiner index and S is the number of species.

The Margalef index (O) was calculated as

D=(Sl)/inN eqn- (3) where S is the number of species and N is the number of individuals.

The Berger-Parker dominanw index (d) was calculated from the equation:

d=N"JNt ecin (4) where N, is the number of the most abundant species and N, is the total number of species. In order to ensum that the index inmases with inmasing diversity, the niciprocal fom of the measun was used (Magurran 1988).

The Q statistic (Q), which is a measun of the interquartik slope of the cumulative species abundance curve, was calculated first by choosing the upper and lower quarliles, (i.e., the points below which 25% and 75% of the species abundances lie) (Kempton & Taylor 1974). Once the quartiles are seleded. the Q statistic was calwlated by:

Q=(0.5nR, +Zn, + 0.5n,)/log2(R2/R 1) eqn. (5) where OSn,, is half the number of species in the class in which the lower quartile falls,

Zn, is the total number of species between quartiles, 0.5n, is half the number of species in the class in which the upper quarüle falls, and Ri and R2 are the number of individuals in the classes with the lower and upper quartiles respecüvely.

The alpha statistic (a) was calculated by:

a=N(1-x)/x eqn.(6) where x is detemined fmm S/N=[(l-x)lx][-ln(1-x)], S is the total number of species and N is the total number of individuals.

The above diveMy indices are measures of alpha divenity which is the divenity of a biological community within a habitat (Southvood 1978, Magurran 1988). Beta diversity is a cornparison of species composition between different communities

(Southwood 1978; Magurran 1988). To assess beta divenity, the Jaccard index (J) was used (Southwood 1978):

J=SJ(S,+Sb-Se) eqnv (7) where S, represents the number of species shared by community a and community b, and S, and Sb am the number of species found in community a and b, respectively.

Dmndognms bared on the Jaccard's index were prepared for carabid assemblages

between, and within, tfeatmenb (Magurmn 1988).

Statistical Analv sis

Data were collected on the 18 fields (repliates) within four treatments

(fams). The treatrnents represented 3 different tillage types (no tillage, medium, or

high). Data from the two high tillage treatments consisted of two farms which were

treated separatey because one of the fans was added a year into the expetiment.

Each replicate was subjed to a particutar trap type (single or barrier), and crop (corn,

soy, or wheat) whem the uop type rotated year to year. Data were examined and the

data wem found to be approximately nonnally distributed. Repeated measures analysis

(RMA) of seven diversity indices (me above) were taken weekly for 9 weeks (see above)

on each feplicate at each position dong a ûansect ove?aie coune of 4 yean (1 994-

1907). The SAS proceâun PROC MlXED (Littell et al. 1096) was used to fit a mixed

mode1 where the effect of repliates within tmatments was treated as a random effed,

while the other fadon, trentment, uop, 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 effed of the fixed factors (treatments) Hile individuel t-tests were used to reveal differences in the least square means of the levels of each treatment.

In addition to examining diversity index differences between treatments, RMA was perlonned on catabid abundanca and richness data wllected 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, aop, trap type). These slopes were subsequently analyzed by titting the model explained above. This analysis was repeated using the slopes fmm 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 my study with those on other carabid populations, published data sets from various disturbance studies were gathered for meta-analysis.

Disturbances in those studies include: dear cutting (Sustek 1981; Lenski 1B82), pollution exposure (Critchley 1972; Freitag et al. 1973; Jamiik 1983; Hejkal 1985; Asteraki et a/.

1992), fotest fragmentation (Niemela el el. 1992, Halme & Niemela 1993) and agncultum (House & All 1981). The rame species divenity indices as used for carabids in my study vuen applied to those data. General Findinas

Thirty genera, and 102 species, of cambids were identified frorn the 20,495 carabid beeties collected during this study (see Appendix 3 for data). Over four yean of collection (1994-1 997) sixty-eight spedes were found on no tillage treatment (NT), fifty- seven on the chisel plowed treatment (MT), and forty-ho and fifty-three on the mold board treatment (HTI and HT2, respectively) (see Appendix 3). Figure 2-2a-l illustrates the abundance of carabid species on each treatment. Species which were found on al1 four treatments include Anisodactv,~~senctaec~sis Fabricus, Bembidion inaequale Say,

Bembidion tetmcolum Say, Bembidion quadrimeculatum oppositutn Say, Chlaenius tricolor Dejean, Clivina fossor LinnB, Harpelus henbivagus Say, Plemstichus melanarius

Illiger, Stenolophus comma Fabricius and Tachys incutvus Say. Agonum muelleti

Herbst occurred fiequently on al1 but NT replicates (Figure 2-2a). Amam 8onelli species were not collected from HT1, but were found on the other treatments (Figure 2-2 b).

Cicindela and Harpelus species were far more common on NT Vian the other treatments

(Figures 2-2 d and h). Dyschirw species were not found on NT or the neighbouring

HTI, but were collected in low numbers fmm HT2 and MT replicates (Figure 2-2 g).

There wem yearly difierences in aie ranked ordei of the most numerous species maiin each tr~atment,but the common species within each treatment were present throughout the study. For HTI nplicates, species which were cornmon throughout the study peflod were Cliwha fossor, Bembidion quadnmculatum oppositum, PIemstichus Fig. 2-2. Pooled abundance of species found on each treatment. HTI and HT2 are the rnold board treatments, MT is the chisel plowed treatment and NT is the no tiII treatrnent.

2-2a. Agonum species s Fig. 2-2. cont. Pooled abundance of species found on each treatment. HTl and HT2 are the mold board treatments, MT is the chisel plowed treatment and NT is the no till treatment.

23c Anisodactylus species

T2 treatment Fig. 2-2 cont. Pooled abundance of species found on each treatment. HTI and HT2 are the mold board treatments, MT is the chisel plowed treatment and NT is the no till treatment.

T

treatment Fig. 2-2 cont. Pooied abundance of species found on each treatment. HTI and HT2 are the mold board treatments, MT is the chisel plowed treatment and NT is the no till treatment.

treatment Çig. 2-2. cont. Pooled abundance of species found on each treatment. HTI and HT2 are the mold board treatments, MT is the chisel plowed treatment and NT is the no till treatrnent.

2-2i Lodceme to Poecilus species

2-21 Pte10S1/chw species Fig. 2-2 cont. Pooled abundance of species found on each treatment. HT1 and HT2 are the mold board treatments, MT is the chisel plowed treatment and NT is the no till treatment. 2-2k Selonophonrs to Shnolophus specks

2-21 Syntomus totmhus species mlanuriius, Tachys imutws. For HT2 replicates, the most common species include

Agonum muel/en, Ptemstichus melenarius, Bembidion quadtiineculatum oppositum. For

MT replicates, the most common species wem Ptemstichus melanarius, and Bembidion quadrimeculatum oppositum. For NT tepliates, the most common species wem

Harpahrs pensyhnicus de Geer, Remstichus melanarius, Bembidion quedrimeculatum oppitum and CWndeIa îùmagenemsa Dejean.

Exotic species wtiich were collected from al1 treatments were: Agonum muellefi,

Amen aenea, Bembidion telrecolum, Bembidion quadrimaculetum oppositum, China fossor, Hatpalus allnis, Harpalus puncticeps, Ptemstichus melanarius, Tmhus discus and Tmhus qwdrlsttfatus (se8 Spence 1990 for list of exotic spedes in Canada). The exotic species Bembidion obtusum Seiv. was found only on HT1 and HT2, and Amara familiafi DR which was collecteci only from NT. The total percent abundance of exotic carabids collected from each treatment are: 42.31 1 in HTl , 77.48% in HT2, 60.59% in

MT and 35.87% in NT. The high percentage abundance is because some of the most commonly cdlected species (i. e. , Agonum mueNe ii, Bembidion quadfimaculatum, Clivina fossor. Rwostichus mlanatius) are exotics. The ratio of exotic species to native species wllected fmm each treatment was 9:33 in Hf1, 10:43 in HT2, 8:49 in MT and 1058 in

NT. Null hypothesis that exotic species are more cornmon in treatments with less tillage can not be rejected Mencontingency tables are used ($ = 1.38, df= 3, p0.01). Çanbid Richness and Abundanw over transects.

For single trap transects, the results of RMA on carabid abundance reveal that there were no significant effects of fam, year or uop type per trap, for trap positions or

üme (F values fanged from 0.56 to 1.84, df ranged from 1-3 and 10-12). For carabid richness over time there was no overall signifiant effect for farms (F-2.33,df=3, 12, p0.12). However, there was a significant year by crop interaction (F=9.03, df=2, 10, p-0.01). Least square means reveal wheat had a significantly different effect than did corn and soy (t=3.03, df=lO, p <0.01, and t=4.17 df=lO, p<<0.01, respectively) but that corn and soy did not differ fmm one another.

For bamer traps, the results of RMA on carabid nchness 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 1-3 and 1-93). 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. 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 1-3 and 1-5). However, there was a significant year by Ume (i.e., collection period) interaction for abundance. Least square means revealed that NT replicates weie significantly different from the other 3 treatments in 1997 when more beetles were trapped in barfier traps in NT than the other treatments.

Generally, then, was no differenw in tmp position or over collection time for richness or abundanœ of carabids collected in fields of the four treatments. Therefore, it is possible to pool data from the traps over time to calculate divenity indices for each treatment.

Jaccard's Similaritv Index

Table 2-3 provides Jaccard's similarity indices for all replicates. The mean similarity within each treatment is: 0.37 for HTI, 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 wem trapped in transects with barrier traps than in transects with single traps. However, species composition found

in the two trap types were not significantly different. Dendograms showed that species composition in NT and MT are more similar to one another than HT1 and HT2. Within treatments, species tend to be most similar between yean , but crop type does not seem to have an effect (Figure 2-3). Table 2-3. Jaccard's index of similarity. 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 (Le., medium tillage) treatment in Waterloo County. NT is the no üllage treatment in Oxford County. The number after the treatment name refen to replicate field. a) between treatments in 1994; b) between treatments in 1995; c) between treatments in 1996; d) behiveen treatments in 1997. Fig. 2-3. Dendograms of carabid assemblages based on Jaccard's index. HTI represents mold board tillage (Le., high tillage) treatment in Oxlord County and HT2 mpmsents mold board Mage treatment in Wellington County. MT is the chisel plowed (i.e.. medium tillage) treatment in Watedoo County. NT is the no tillage treatment in Oxford County. Tho numbrr after the tmatment namo refen to replicate field. wwheat, s=soy, ocom, 94= 1994,95=1995,96=1998,97=1997

a) replicate data pooled for each treatment

Percent Similarity

b) HTl data Percent Sirnilarity Fig. 2-3 cont. Dendograms of carabid assemblages based on Jaccard's index. c) HT2 data Percent Similarity

d) MT data Percent Similarity

e) NT data Percent Similafity Diversitv Indices

The ovenll effect of treatment, crop, or transect. on divenity indices (Q-statistic,

Margalef, richnesr, evenness, Shannon-Wiener, alpha and Berger-Parker) was not significant. Hovuever, there were some year by treatment interactions. Thus it is necessary to look at the simple effects of treatment by year cornparisons with least squares rneans. For the Q-statistic, in 1994, HT2 was significantly different from MT and

NT (k2.85, df=28, p4.01, b2.10 dt28, peO.05, respectively), end, in 1998, HTl was significantly different fiom NT (t=2.11, df=26, p<0.04). For richness, in 1gm, NT war significantly diffant from HT1 and MT (t-2.14, dfn28, p<0.04, e2.55 df=26, p

For Berger-Parker, in 1995, NT was significantly different from HT2 (1-4.12, df=26, p<<0.03), in 1996, HTI was significantly diffennt from MT (t=3.32, df=26, p-0.03) and, in 1997, NT was significantly different from MT (t=2.89,df=26, peO.01). For alpha, in

1994, HT2 was significantly different from MT (t=2.89, df=26, p

Carabid species diversity was calculated for 45 published carabid data sets hm

10 studies (Table 2-4). No one diversity measun, was better than aie others et pinpointing the sites that wre disturbed. For example, Shannon-Weiner (H') measures for Critchley's (1972) data indicated that the site with greatest exposure to pesticide spraying had greater divenity than the control site. Meanwhile, Margalefs index suggested the opposite to be true. Although Lenski's (1982) data shows that H' was greater in forested areas than in clear cuts, Margalef, Q-statistic and alpha did not. Table 2-4. Diversity indices for published carabid data. R is nchness, A is Abundance, H is the Shannon-Wiener index, E is the Shannon evenness index, BP is the Berger- Parker index, Al is the alpha divenity index, M is the Margalef index, Q is the Q-statistic. For descriptions of the sites see studies.

Rtfrnnco sito RAH'E BP Nkmrli et al. (1992) upknd ispm fores! 18 734 1.39 0.49 0.57 aspen popîar fmt 21 403 1.96 0.64 0.35

Halme and Niemafa (1 993) contiguous forest large fragment medium fragment srnall fragment we surroundings

Sustek (1 981)

Frietag (1 973)

Critchley (1 972)

lenski (1 982) mit forest mis clclrcut et foreSI er dlaricut min forest rmn ckucut

Homand All(t981) corivaritlonrl mm fescua lkld woods DISCUSSION

In my study some carabid species were cornmon in al1 treatments. Those species include: Anisodactylus sanctaec~sis,Bembidion inaequele, Bembidion tetracoIum,

Bembidion quadrim%cuI8tum, CMmnius tricolor. cIiMn8 fossor, Harpelus hehivagus,

Plemstichus mdanartus, Stenolophus comma and Tachys incurvus. The occurrence of exotic species was not higher in more disturbed treatments than in the less disturbed treatrnents. The exotic species I collected are well established within the carabid assemblages (Spence 1990). Those species have been part of the carabid fauna of

Canada for a minimum of 30-40 years (Lindroth 1961-1QW), Le., at hast 30-40 generationr, and an, unlikely to act as colonking exotic species which occur in higher abundance than native species in disturbed environments (Elton 1958).

I used pitfall traps to collect carabids for my study. Although there has been some discussion about the reliability of using pitfalls to capture carabid beetles (e.g., Digweed et al. 1995; Morrill 1%O), then have been studies (e.g., DuMne & Legrendre 1997, Kromp 1999) which indicate that pitfall trapping is a reliable method of collection. Because the species collected in the barrier baps were similar to single traps, it is assumed Viat a representaüvo sample of carabid species from each fann was collected with single trap transects. If the single trap transects did not provide a representative sample, then the species compositions between trap types would have been different. Although the major diffemnca arnong treatments was tillage practice, my nuIl hypothesis, that there would be no change in divenity with increasing tillage, could not be rejected for any of the divenity indices used. Several studies have shown that carabid divenity was greater in fields under minimum tillage (Dritschillo (L Wanner 1980;

House & Alllg8l; House & Stinner 1983; Ferguson & McPhenon 1985; Bnist et al.

19û6a; Weiss et al. 1990; CBrcarno 1995). However, other studies either indicate tillage does not affect carabids (Tyler & Ellis 1979; Dritschillo 4% Etwin 1982; Hokkanen 8

HoIopainen 1986; Krornp 1989; Mack & Buckman 1990;Tonhasca 1993) or that carabids are more abundant in wnventionally tilled fields (Edwards 1979). Thus. my study joins studies which suggest that carabid beetle divenity is not affected by tillage. Unlike these studies, which mainly used only Shannon-Weiner and species richness, I took a novel approach and examined a range of different diversity indices, each with different biases and methods of calculation. I wanted to determine if there is decreased dominance

(Berger-Parker), increased evenness ( Shannon evenness), or increased richness (alpha,

Q-statistic, species richness. Margalef, Shannon-Weiner) with less tillage. Even though I warnined a range of divenity indices I found that no one index provided a consistent indication that carabid divenity was different between cmps or farms.

The present results ansupported by the results of the meta-analysis of published carabid data sets in which none of the different dkenity indices could predict disturbance any mors reliably thon any of the othen. Again, the nuIl hypothesis that there would be no inmase in divenity wiai disturbance could not be rejeded. This suggests that diversity indices do not provide reliable tools for detecting disturbance. Indices are commonly used because ecdogists consider them as a standard approach to addressing questions of disturbance. Just because they are commonly used does not, however, metan that thsy am mliable. The indices which are usetd Vary fmm study to study, and pemaps the results which researchen use are aie ones with which the nuIl hypothesis cm be rejected (Peters 1991).

Alternative methods to measure disturbance are based on distributions of carabid beetle size, and species divenity and abundance. In chapten 3 and 4, the nuIl hypothesis that there would be no deviation from log normal distribution of carabid size

(Chapter 3), and species divenity and abundance (Chapter 4), with increasing disturbance which is represented by tillage are examined. CHAPTER 3: THE EFFECT OF TILLAGE DISTURBANCE ON BODY SUE

DISTRIBUTIONS WITHIN ASSEMBLAGES OF CARABIOAE

In this chapter I examinrd the novel hypotheses that there would be decreasing divesity of beetle species sizes, imnasing mean bretle size, and increasing deviation away from log normal distributions of size, with increasing disturbance as represented by tillage. Because the nuIl hypotheses were not rejected, it is possible to wnclude that tillage encountered in this study does not have an effect on carabid community structure.

This conclusion is the same as that in Chapter 2 in which divenity indices generally indicate carabid beetles were not affected by tillage. The log normal distributions of size suggest carabid beetle assemblages form a fundional gmup.

INTRODUCTION

Although much work has been done on species diversity, and species divenity and abundance models, less attention has been paid to species size distributions within hrncüonal groups as desaibed in Chapter 1 (May 1975; Griffiths 1986). Species' size distributions am often polymodal (see Griffiths 1986). Thmo commonly offered explanations for this pattern an: 1) nsoum utilization by predatom reflects polymodal size distributions of prey (GriffiVis 1986), 2) cornpetition results in the divergence, not convergence, of species' sizes (Fretwell 1978; Kirchner et a/. 198O), and 3) the sizes of predaton depends mosUy on the existence of a size dass of smaller organisrns as a food source (Popova 1978; Kerr 1979). However, many of the studies on which these explanations anbased have used species colledions which reflect guilds or communities (see Chapter 1) not funcüonal groups (for exemple African bovids; Jarman

1874; fmsh water fish; Popova 1978; nematodes; Roff 1981).

The size distributions of species within funcüonal groups should be log normal because in undisturbed habitats the range and variability of energy and resources are log normally distributed (Harvey & Godfray 1987). Theoretically, there should be few very large and very small species, whereas most species would be of intermediate size.

Similady, there would be some resources with little energy, few with much energy but most resources would have intermediate energy. Log normal distributions of species' body size thus can be seen to reflect how energy and other limiting resources are allocated within functional groups (Brown & Maurer 1986; Siemann et al. 1996). By analogy, an adaptation to Sugihara's (1980) arguments on !og nomal distributions of divenity and abundance is that the energy and resounes within a niche should be split by species within a functional grwp. Thus' the fractions, within a niche, of the energy and resources that are apportioned to any given species should be proportional to that species' body size. Hence, a fourth explanation for polymodal distributions of species' body sizes is that they represent the existence of a number of functional groups (one per mode) eacfi with its own cornpetitive interactions.

In undisturbed habitats, body size distributions of speues in a functional group should be log normal (Sugihara 1980; Preston 1980), but once a habitat is disturbed, it has been argued that then should be more larger species than smaller species because larger species are able to tolerste a greater range of environmental conditions

(see Griffiths 1986). However, disturbed habitats tend to be populated with colonizing species which are usually r-strategists (Hutchinson 1978). R-strategists tend to be small and often are vagile (Hutchinson 1978). Disturbed habitats may also have lower divenity of species' sizes within fundional groups in situations where disturbance decreases the number of microhabitats (Lomau 1988 on Carabidae). Because large species utilize a disproportionately large share of the resources within local ewsysterns (Strong et ai.

1979; Brown & Maurer 19W) they are usually dominant in interspecific competition and exdude smaller species (Morse 1980). This has been show for bitds (Brown & Maurer

1988); fish (Smith a Taylor 1972); mdents (Brown & Munger 1985) and plants (Turner &

Brown 7982). As far as I andetemine, there are no data on cornpetitive interactions

between large and small Carabidae. In fact, arguments conflict as to the general

importance of competition within carabid assemblages (see Thiele 1977; Loreau 1988).

The distribution of species' body sizes may become either increasingly bi-modal or log series with increasing levels of disturbance. A log series distribution would resuît if

large species out-compete, through domination, almost al1 the small species for limited

resources. Bi-modal distributions would occur if the smallest species were able to out-

compte large species in niche spaces which are small, and not exploitable by large

species in a manner analogous to the eWof refugia for prey in predator prey

population dynamics (Andnmtha & Birch 1954). FU~~WIIIO~~ it rnay be energetically

costly for large rprdes (espdally pmdaton) to feed on food items below a certain sire

(Pearson & Mury 1979). Thus, those food items would be left available for small species. Published examples to support aie ideas introduced above, with respect to the effects of disturbance, are rot avaikible because this approach has not been explored.

Thus, my investigation of species' body size distribution as used to assess the

hypothesis of deviation of log nonnality in parallel with examining species diversity and abundance (Chapter 4) is novel.

In this chapter, the influence of a gradient of disturbance on carabid beetle size

distributions is investigated. Agricultural fields with no tjllage are less disturbed than are

fields with chisel plowed or mold board plowed Mage. lncreasing tillage has been shown

to decrease the number of micro habitats available to the beetles (Stinner et al. 1988;

House & Pannelee 1985). Thus, on fams (Le., treatments, see Chapter 2) which are

npnsented by more tillage, the body size distributions of beetles should deviate

increasingly away from log nomality. Body sizes are examined both by using mean lengths and the approximate species' volumes (Le., biovolumes, see below) . The

divenity of species' sizes should decrease, and large species should alto be found

disproporüonateiy mon frequently, on treatments represented by high Mage if the

argument of Gnffiths (1986) is accepted. Notwithstanding that I did not find divetsity

indices to be useful, t examined the divenity of carabid sizes by using a modification of

the Shannon-Wiener index. I modified this index rather than one of the other ones

because it is based on information theory (Magurran 1988). ln this case species' lengths

were considered rather than taxonomie classifications of species which were used in

Chapter 2. METHODS (For site descriptions and collection methods see Chapter 2.)

Shannon-Wiener divenity measures were applied to body lengths of the species found in each mplicate (i.e., field) of evrry treatment (Le., fam). In this case

Hldi- = 4( p,lnp, ) eqn. (1) whete p,=L)L. 4 is the length of the ith spedes in the samplr and L is the sum of the lengths of al1 carabid species found in the sprdfic collection in question. Categorization of species lengths was made by mean species lengths as reported in Lindroth (1961-

1969).

In addition to H'diwni

(L,.)collected for each replicate on the 4 treatrnents was calculated. Repeated measures analysis (see Chapter 2) was peffonned for H8- and La,,.

The distributions of carabid lengths and carabid biovolumes for each treatment were graphed. The biovolume of carabid species was detemined by rnultiplying the mean length of beetle species by the mean width and by the mean depth according to the methods of Siemann et al. (1999). Because carabid body shapes (for the species colledecl in this study) are similar, the proportionate measures of width and depth in comparison to length was detenined by measuring the width and depth for 10 beetles from 10 different species in my collections (Table 3-1). The correlation coefficient is high for both carabid lengths and width (+ =0.94), and Iength and depth (?= 0.90) (Figure 3-

1). Thus. it is possible to use the proportions between length and width, and length and depth. to approximate carabid biovolume. The Mdth of canbids is on average 0.42 times the length, and the depth is on average 0.27 times the length. Although biovolume as calculated is just an approximation of mal volume, it is suificient for my study. Moreover, pmcise measurements are not needed because the distributions of biovolumes were converted into octaves for furthet analysis. The octaves (or biovolume size classes) were are follows: species which are 1-2mm3 were put into dass 1, species which are 2.1-

4mm3 were put into dass 2, species which are 4.1-8mm3 were put into class 3 and so forth. The distributions of lengths were not put into octaves because the range of lengths was not suifident to be broken into meaningful octaves. The data was analyses for their

similatity to kgnonnality by the Lilliefon diagnostic (Sokal & Rohlf 1995) by which the probability of rejecüng the nuIl hypothesis was used to increase the probability of rejeding the nuIl hypothesis. Table 3-1. Width and depth measurements for 10 beetles of 70 different representative species. Standard deviations are in parentheses. -

SPECIES length (mm) depth width Agonum cupripenne 8.40 (1.47) 2.48 (0.27) 3.45 (0.09) Agonum muellen 8.35 (0.62) 2.30 (0.30) 3.40 (0.25) Bembidion inaequale 5.10 (0.06) 1.28 (0.02) 2.18 (0.04) Bembidion quadrimaculatum 3.25 (0.05) 0.90 (0.05) 1.58 (0.05) Bernbidion tetracolum 5.50 (0.50) 1.43 (0.03) 2.29 (0.05) Cicindela fornosa generosa 17.00 (1.94) 5.48 (1.21) 6.68 (0.98) Clivina fossor 6.00 (1.21) 1.î8 (0.08) 1.76 (0.06) Harpalus pensylvanicus 12.65 (1.45) 4.50 (0.32) 6.00 (1.50) Loricefa pilicomis 7.75 (0.70) 1.80 (0.58) 3.60 (0.43) Plerostichus melananus 15.50 (1.20) 4.50 ( 0.35) 6.15 (1.15)

Fig. 3-1. Measures of a) depth vs. length and b) width vs. length for 10 beetles from 10 carabid species (see Table 3-1, and text). The correlation coefficient is high for both length and depth (r'=0.90),'and carabid lengths and width (9=0.94).

a) mean depth (mm) vs. mean length (mm) b) mean width (mm) vs. mean length

length (mm) lenoth (mm) RESULTS

The mean sizes of species colleded in my study ranged in mean length from 2.25 mm (Tachys incurws Say) to 22.80 mm, (Harpelus c~lbiinosusFabricius). Figun 3-2 shows the Iength distribution of carabids that were wllecîed on each treatment (Le., fam). On al1 treatments the distribution of MeUe species Iengths was log normal. On the treatrnents which are repmented by high Mage 2 = 0.77, df = 6, p = 0.60 for HTI ,

2 = 0.55, df = 5, p > 0.90 for HT2, on the treatment represented by medium tillage MT $

= 0.53, df = 5, p > 0.90 and on the treatment represented by no Mage x2= 0.53, df = 5. p>0.90.

Figure 3-3 shows the biovolume distribution of carabids that were collected on each treatment. On al1 treatments the distribution of biovolumes was log normal. On the treatments which were represented by high tillage $ = 0.90, df = 8, p = 0.25 for HTl;

)(? = 0.92, df = 8, p = 0.40 for HT2; on the treatment represented by medium tillage MT x2 = 0.93, df = 8, p = 0.42, and on the treatment repmsentad by no tillage 2 = 0.88. df = 8, p = 0.20.

Table 3-2 lists divemity of lengths, and mean Iength, of the carabid species collected within each treatment With npeated measures analyses for divenity of carabid species sites (Hmdluriïr) there was no signifiant effect (p>O.OS) of treatment, tnp type, year or crop (F ianged from 0.50 to 2.03). For mean length, there was no significant effect (p>0.05) of trap type, year or crop (F ranged from 0.38 to 3.22),but the tmatmetnt effect is significant (Fs25.98, df=3. p<

Fig. 3-3. Carabid species biovolume by volume class from poolad replicate data for treatrnents representing high Mage (HTI and HTZ), medium tillage (MT) and no tillage (NT). Volume classes: 1 is1.1 -2.0mm3,2is2.1 -4.0mm3,3is4.1 -8.0mm3,4is8.1 -16.0mm3,5is16.1 -32.0mm3, 6 is 32.1 - 64.0 mm3, 7 is 64.1 - 128.0 mm" 8 is 128.1 - 256.0 mm3, 9 is 256.1 - 512.0 mm3, 10 is 512.1 - 1024.0 mm5and 11 is 1024.1 - 2046.0 mm3.

12345678910tl voluma clam

Fig. 3-4. Mean carabid species length cdlected hom treatments in each year representing high tillage (HT1 and HTZ), medium Ullage (MT) and no tillage (NT). beetles were wllected from the treatments, the treatment npresented by no-tillage was always significantly different from al1 other ûeatments (Figure 34). The treatment npresented by chisel plmdtillage was si~nificantlydiffemnt from one of the treatments which mpresent mold board Mage (HTI, see chapter 2) but only in 1996.

DISCUSSION

The nuIl hypotheses tested in this chapter are that there would be no difference in divenity of beetle species' body sizes, no diffetmnœ in mean beetle length, and no differences in deviation away from log normal distributions of size, with incteasing tillage distuhance. The nuIl hypothesis of a log nonnal distribution of size cannot be rejected.

The distributions of biovolumes and lengths were log nomally distributed, and did not deviate with incteasing degrees of tillage disturbance. The nuIl hypothesis for beetle length was rejected because the mean length of carabids in the treatment representing no Mage was diffemnt, in fad greater not less, than the mean lengths, in the treatments representing more tillage. According to Griffiths' (1986) argument, it would be expected that there would be disproportionately more large spscies than smaller species on higher tillage fields because larger species are able to toterate a greater range of environmental conditions. These results rnay have occurred because either tillage does not provide elctrerne disturbance for carabid beetles or, in other words, what we perceive as greater eQemrs are not extrema$ to other organisms.

Cornpetition for resowees can be reflected in log nomal distributions of size in fundional groups (Brown & Maurer 1986; Siemann et al. 1996). The log nonnal size distributions in the carabid beetles populations I wllected suggest that carabid beetles ana functional group and competition exists between beetle species (Brown & Maurer 1986; Siemann et al. 1996). This conclusion about intenpecific competition supports Loreau (1989) who suggested that intenpecific cornpetition occuts between carabids, but is conûary to the studies by Thiele (1977) and den Boer (1985) who stated that there was no intenpecific competition. One differencs between my study and the othenr cited is that they diredly compared interactions between species whereas I examined the size distributions which are proposed to be the outcome of campetition. A possible benefit of my approach is that the overall effect of competition is reflected, not just the competition between sets of species which are assumed by the researcher to competitively exclude one another (e.g., Remstichus vulgaris, a field species and in competition with Abax ale4 a forest species under Iab conditions (Thide1977)).

The novel hypotheses tested in this chapter are that there would be decreasing diversity of beetle species sites, increasing mean beetle size and increasing deviation away from log nomal distributions of size, with increasing disturbance as represented by tillage. Because Vie hypotheses were not accepted, it is possible to conclude that the tillage encountered in my study does not have an effect on carabid community structure. This finding supports the conclusion found in Chapter 2 in which divenity indices generally indicate carabid beetles wem not affected by tillage. Species diversity indices, and size distributions are two approaches for detecting the effect of disturbance on functional groups. A third approach is to investigete log normal distributions of divenity and abundance of species, and detemine if there is increasing deviation from log nonnality with grnater disturbance. This approach is the subject of Chapter 4. CHAPTER 4: CAMBIO BEETLE ASSEMBLAGES AND THEiR FIT TO DlVERSlTY AND ABUNDANCE MOOELS UNDER A GRADIENT OF TILLAGE OISTURBANCE

ABSTRACT

I examined the hypothesis that carabid beetles are a functional group by using distribution of species diversity and abundance. The resuits of Chapter 3 suggested that carabid beetle assemblages are a functional group because they have log normal size distributions. However, those results might merely be statistical artifacts. In this chapter, I suggest that carabid beetles am not a functional group because they are not represented by log normal distributions of species divenity and abundance. I found that carabid populations were al1 geometrically distributed in al1 treatments (Le., fams which represent different degrees of tillage). I also examined the hypothesis that carabid populations would deviate increasingly from log nomality with increasing Mage disturbance as represented by different treatments. Because the beetles were never represented by log normal distributions they did not deviate away frorn log nonnality with increasing distufbance. These conclusions wen compared to those from a meta- analysis of forty five published carabid data sets which confimis that carabid beetles are not represented by log nomality under undisturbed situations INTRODUCTION

Studies on aie eflect of tillage on carabid divenity have not been conclusive (see rekrences in Chapten 1 and 2). In the treatments (i.e., fams which represent different degrees of tillage, see Chapter 2) used for this study, carabid beetles do not appear to be affected by tillage disturbance, as assessed by either dhersity indices, or deviation away from log nomal distributions of body size (se8 Chapten 2 and 3). Although using deviation frorn log normal distribution of riz0 is a relatively mvel approach to studying the effect of disturbance on a functional group (but see Preston 1980). use of diversity indices is standard ecological practice (Magurran 1988). However. despite the common usage of divenity indices by ecologists, they do not always detect disturbance. In Kevan et d.'s (1997a) study on pollinating bee populations, divenity indices did not reveal any effect of pesticide disturbance whereas there was deviation from log normal distributions of divenity and abundance. Thus, another approach for studying the effect of disturbance on a functional group is to investigate whether or not the species diversity and abundance distribution of the group in question deviates from log normality. This approach has been used with pollinating bees (Kevan et al. 1997a), Lepidoptera (Laroca et al. 1989; Hill et al. 1995) and soi1 arüiropods (Higvar 1994), but has not been used with carabid beetles.

The fint hypothesis to be tested in this chapter is that carabid beetles (assumed to be a fundional group on the basis of size distributions; see Chapter 3) conforni to log normal distributions of species divenity and abundance. In this chapter, I have used the approach of exarnining the fit of log normal distributions of divenity and abundance to address the question of whether uirobid beetles can be considered as a functional group when canbids anconsided taxonomically. Because species within funcüonal groups compete for resources, detennining whether of not carabid beetles belong ta a functional

67 gmup is useful for addressing the question of competition between carabid species.

According to sorne researchen (Loreau 1988 ) there is intenpecific competition between carabids, whemas oüwn (Thiele 1977; den Boer 1980) suggest there is not (see also Chapten 1 and 3). By examining the distribution of species diversity and abundance of carabid species it is possible to ask if competition between species occun from a whole grwp (community) perspective mthet than by examining individual species' interactions. The benefit of this approach is that the entire carabid assemblage is taken into anount, rather than just a few memben as in the more restricted studies of Thiele (1977). den Boer (1980) and Loreau (1988).

Another hypothesis tested in this chapter is that carabid populations increasingly deviate from log normality with increasing degree of Mage. The results of my study are wmpared to a meta-analysis for species diversity and abundance of for(y five published carabid data sets. By inclusion of those data sets it is possible to determine if the carabid species divenity and abundance patterns found in my study are typical of carabids, and to assess the overell applicability of species deviation of species divenity and abundance from log norrnality under disturbance.

Deviation fmm log normal distributions of diversity and abundance cm be used as indication of disturbance (see Chapter 1). However, in the event that assemblages of carabid beetles did not fit log nomal distributions, fit to log series or geometnc distributions of divenity and abundance can be assessed. By describing the distribution which lits an assemblage of species (even if that assemblage does not constitute a hindional group) it is possiMe to make infennces about some of the cornpetitive intnctions between species. METHOOS (For site descriptions and collection methods see Chapter 2)

Carabid data from each treatment were pooled to ensure that the sample rizes would be suffident to allow for statistically rigorous testing of differences between divenity and abundance models. Thme divenity and abundance models were examined: geometric, log series and log normal.

Magurran (1988) suggests that the simplest method to assess whether or not a geometnc series represents a biological comrnunity is to use rank abundance cuwes.

The log series distribution is represented by:

ax, ( ax2 )12, ( a$ )/3, ... ( axn )ln eqn. (1) ax is the number of species predicted to have one individual, ( a$ )/2 those with two and so forth (Fisher et al. 1943; Poole 1974; May 1975). Magunan (1988) outlines the procedure to obtain a as follows. The total number of species, S, is obtained by adding all the tenns in the series which reduces to: S=a[-In(1-x)] eqn.(2) with x being estimated from:

SIN = (1-x)/x[-ln(1-x)) eqn. (3) where N = the total number of individuals. Because a and N summarize the distribution and are related by: N = aln(l+N/a) eqn. (4) The following equation can be used to obtain a a=[N(-x)x eqn.(5) To detmine if my treatment data sets canfonned to log normal distributions. they wem gmduated to a truncated cuwe udng the following general equation (Preston 1948; Whittaker 1970; Pielou 1969):

69 S(R)=S@xp (-a2*) eqn.(6) Species abundances wem trsnsfomed using log, to generate intervals of octaves. S, is the number of species within the modal abundance octave R,. In eqn. 0, S(R) is the number of species in an octave R octaves from Ro and 1/a is the width of the distribution a=l/((f2a). This constant, a, for a canonicol log nomal relationship ha$ been found to be approximately 0.2 (Hutchinson 1953; Whittaker 1972; Colinvaux 1973).

For species distributions that had modes, log nomality was tested. A crude mode in log normal distributions of divenity and abundance can be seen in histograms as the value which occun 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 Kolmogorov-Smimov test, was used to indicate whether or not my data was log normal. To deted log series distributions, the Kolmogorov-Smimov test was used. As well, rank abundance graphs and frequency distribution histograrns were prepared for pooled carabid data from each treatment.

To compare the tesults of my study with other carabid populations, complet0 published carabid data sets from various disturbance studies were gathered for meta- analysis. Disturbances in those studies indude: clear cutting (Sustek 1981; Lenski 1982), pollution ecpasure (Critchky 1972; Freiteg et al. 1973; Jaroiik 1983; Hejkal 1985; Asteraki et a/. 1992). forest fragmentation (NierneIl et el. 1992; Halme & Niernela 1993) and agriculture (House & All1981). The same species divenity indices as used for carabids in my study were applied to those data (tee Chapter 2). Alsol rank abundance distributions mmexamined to determine if the findings in my study were similar to those from ottiers. RESULTS

The spedes abundance distributions for each hm had long tails of tare species. This is shown in rank abundance graphs (Figura 41) and in frequency distribution histognms (Figure 4-2) of pooled wmbid species data for each treatment. The carabid data did not show a mode in the frsquency distribution histognms, and thus no veil line was presrnt. Although when them is no crude mode, then may be no statistical difference between log series and log normal it is necessary that species with intemediate abundance be distinguishable from rare species for biological significance to the assessed (as per Sugihara 1980). Henwforth for the sake of brevity, I indicate the above conclusion with 'Because there is no mode, the data did not confonnity to log nomality". All the treatments did not conforni statistically to log series distributions: for Hfl 2 = 18.75, df = 9, p< 0.05: for HT2 x2 ~23.30df = IO, p

As wiai the carabid populations in my study, al1 the published carabid species abundance distributions had long tails of rare species. This is shown in rank abundance graphs (Figure 4-3). Because there is no mode, the data did not confonn to log nomality. Table 4-1 lists the log-series analyses for 45 data sets, of which 9 were not log series. In al1 non-log series data sets there were many rare species (Figure 4-3), suggesting that they am geometrically distributed (Magumn 1981). Fig. 4-1. Rank abundance plots of carabid beetles collected in rny 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.

Fig. 4-2. Frequency distributions of carabid species collected frorn treatments representing high tillage (HT1 and HT2), medium tillage (MT) and no tillage (NT), in relation to their abundance: ' Fig. 4-3. Rank abundance plots of carabid beetfes from data sets included in my meta- analysis on a logarithmic scale against the species' rank, in order from the most abundant to the hast abundant species.

Fig. 43a. Data from Niemala et al. (1992)

4upland aspen forest +aspen-poplar forest +spruce bog +lakeside forest +meadow

1 11 21 31 species sequence

Fige44b. Data from Hahne & Niemala (f993) +contiguous 100 -t- large fragment

10 6medium !! fragment f 1 s 4smallfragment t 01 +Edge

species sequence

Fig.4-3~Data from Sustek (1981)

+surrounding -0- clearing

0.01 1 11 spacks sequence 73 Fig. 4-3 cont. Rank abundance plots of carabid beetles from data sets included in my meta-analysis on a logarithmic scale against the species' rank, in order from the most abundant to the least abundant species.

Fig.4-3d. Data from Frietag et a/. (1 973)

'Oo 1 -o-stnl +stn2 +stn3 +stn4 +stn5 +control

0.1 4 l v 1 1i species sequence

Fig. 43e Data from Critchtey (1972)

+control +low dose +high dose

species sequence

Fig. 4-31 Data from Lenski (1982)

+rmsforest +rmsclearcut L_erforest -6- erclearcut -O- rmnforest +rmnclearcut Fig. 4-3 cont. Rank abundance plots of carabid beetles from data sets included in my meta-analysis on a loganthmic scale against the species' rank, in order from the most abundant io the least abundant species.

Fig. 4.3g Data from Jarosik (1983)

3 1O +Cernpolî s +Cernunpofl 2' +Libipoll 7 P +Libiunpoll 0.1

rpecies sequence

Fig. 4.3h Data from House & All(1981) 43- Conventional sOY 4Conservation SOY +Fescue

Fig. 4.3 Data from Hejkal(1985)

10 -O- 1rnanth 8 42years 1 46years 4C s -1)- 17 years CI a 0.1 +22 years Table 4-1. Summary of Chi square analysis for log series distributions for data sets in my meta-analysis. R is richness and A is abundance.

Reference site Niemala et al. (1992) upland aspen fores1 aspen poplar forest spruce bog lakeside forest meadow

Halme and Niemala (1993) contiguous forest large fragment medium fragment small fragment edge surroundings

Sustek (1981)

Frietag (1973) stalion 1 stalion 2 station 3 station 4 slatbn 5 conlrol

Critchley (1972) control low *se high dose

Lenski (1 982) mis forest nns clearcut sr forest srclaarcuf ' nnforest rmn clearcut

Jarosik (1983) œrnpotl cernunpoll libipoll libiunpoll

House and All(1981) conventional soy conservation soy fescue fmld Woods

Hejkai (1985) 1 monlh 2 pan 6 yean 17 years 22 yean

Asteraki et al. (1992) DISCUSSION

The diversity and abundance of carabid beetle assemblages in my study (Figures 4-1 and CZ),and in published studies (Table 41, Figum 43), are either log series or geometrically disûibuteâ. In fact, log normal distribution of divemity and abundance is not repmsented in any carabid assemblaga, even in undisturbed sites (see figures 4-3 for House & AI! 1981; tenski 1982; Niernelb 1992; Halme & Niernela 1993). Thus, carabid beetles cannot be used as indicaton of disturbance when deviation from log nomal distribution of diversity and abundance is used. Moreover, and more interestingly, these findings suggest that carabid beetles do not fom a funcüonal group. This conclusion is in contra& to the results on size distributions presented in Chapter 3.

The distributions of size, as reflected both by biovolume and length, are log normal, suggesting that there is cornpetition, and division of resources, between carabid species (Brown & Maurer 1986; Siemann et al. 1996). However, because of large size diffetences among spedes it is questionabk whether wmpetition occun between many canbid species. Carabid feeding behaviour is body size dependent (Müller 1985), and between carabid species, size similarity is an indirect rneasure of likeness in prey preference (Bnist & House 1988; den Boer 1980). The more than ten fold size difference hom the smallest to largest beetle species suggests strongly that they consume different foods and that they occupy separate portions of the micro habitat (Thiele 1977). Maximum relative prey size tends to inuease the body size. Small species (less than lOmm long) feed only on prey with smaller body sises (Loreau 1988) because they are limited by theif ability to catch, or chew, a number of prey types (Hengeveld 1980). In compafison, large carabid species (longer than 10mm) are not restrided to smaller prey (Loreau 1988). As a result, large carabid species tend to have a Mder prey spectrum than do smaller species (Rushton et al. 1991). Additionally, large species generally use a

77 disproportionately wider share of the nsourœs within local ecosystems (Brown & Mauer 1996; Siemann et al. 1996). Thus, although date for Carabidae are wanting, it is reasonabk to suggest that lsrger carabid species catch more. and a greater divenity, of

PreY

If species within a functional group have overlapping resource requirements, energy would be paitiüoned among species acconling to a log normal distribution (Harvey 8 Lawton 1986; and see Chapter 3). Because there is such a size disparity betwwn carabid species, it is likely they do not partition available resources log nomally, and that there is little intenpecific cornpetition between beetle species which are greatly different in size. Thus, even though by size distribution it might be suggested that carabid beetles am a single functional group, it is more reasonable to conclude that they are not. The log normal distribution of carabid size may merely be a consequence of the Central Limit Theorem.

Species assemblages which are useful as indicators of disturbance, when disturbance is rneasured by deviation away from log nonality, must be a functional gmup (see Chapter 1). Carabid beetles, it appean, are not useful as indicaton of disturbance because they annot a single fundional group. There is likely to be little intenpecific ampetition between carabids because of their size differences (see above), but also because carabid feeding behavior is diverse (Larochelle 1990; Appendix 2). With the exception of a few specialists, most carabid species are opportunistic feeden even though they are often considemd as pndaton (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 divene, between larvae there are additional differences in feeding habits (Lindroth 196 1-1989; Thiele 1 977; also see Appendix 1). Because of the diversity of feeding behaviours in the different life stages of the beetîes, distuhances would not have been expected to have a uniforni effect on al1 species. Because species as adults and Iarvae have different feeding behavioun, there is not netessarily direct cornpetition within life stages, which in tum suggests that there may not be cornpetition between al1 of the carabid species.

Because wnbid bkisas r gmup have feeding behavion 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 detemined groups (i.e., small, medium and large) so that aiere would be increased likelihood that cornpetition occun within a designated group. To become a complete functional group, each of those sub- groups rnust be united with additional species from other taxa. For example, carabid beetles together with staphylinid beetles of similer sizes and feeding behavioun may rnake a functional group which can be analyzed for log normality. Unfortunately this idea is very diffïcult to test because staphylinid beetles are limited by taxonomie factors and the ecology of many of the species are not well documented.

Log series and geometric distributions are usually considered to result from niche preemption (Magunan 1988; Thomas & Shattock 1986; May 1975; Boswell & Patil 1971). However, for carabids this argument cannot be invoked. If niche pre-emption occurred, the size of the species which dominate from one year to the next would be consistent. In this study, and in others (see references in Table 4-1). the size of carabid species which dominate from one year to year was not consistent (see Chapter 2 and Appendix 3). If the carabid assemblage is made up of a number of incomplete fundional gmups, then a log series distribution is to be expected. Within each size limited sub- gmup there would be changes in dominance, and carabids would not necessarily be the taon to which the dominant species would belong. By haphazardly categorizing memben from different incomplete functional groups together, a log series, or geometric, distribution would be expected. Canbids are not effective as indicaton for a detailed undentanding of the dynamics of disturbance within ecosystems because they are not a funcüonal group. My findings that carabid beetles are not useful as disturbance indicators are important because they lead to hypotheses about the type of organisms which are useful. Communities which should be targeted for providing potential standards (see Chapter 1) based on divenity and abundance models should include species which feed on a single type of food, species which da not have size dependant feeding, and species which are more or less a complete fundional group. Chapter 5 examines more closely some of these assumptions with published data on a variety of invertebrate communities under different disturbance regimes.

(e.g., Freitag et al. 1973; Thiele 1977; House & All 1981; Hance & Gregoire-Wibo 1987; Fan et a/. 1993; C4namo 1995) it is important to move beyond them and explore other possibh fundional groups for their potential. Data sets fmm a range of organisms (induding spiders, mawlepidoptera, soi1 mites, bees and diving beetles) under different disturbances (ranging frorn clear cutting, industriel pollution and agricultural activities) are explored in this chapter to further examine the hypothesis that fundional groups deviate from a log nomal distribution of divenity and abundance when exposed to disturbance.

MATERIALS AND MET HODS

To compare the results of carabid species assemblage structure with those of other invertebrate species, complet8 published data sets from disturbance studies were gathered. One difficulty was finding published data sets which had complet8 species lists (e.g., SekuliC 1987; Weiss et al. 1990). Many papers which include data sets do not report species that ocair in low numben. Complete data sets in studies of disturbance were found foi: logging (spiders: Coyle 1981), physical disruptions (diving beetles: Nilsson & Soderberg 1998; soi1 mites: Kevan et al. 1995; spiders: Haskins & Shaddy 1996; Doane & Doane 1979; bees: MacKay & Knerer 1979) and pesticide exposure (Lepidoptera: Butler et al. 1995). I examined the species diversity and abundance distributions in those data sets (see Chapter 4 for methods). I also evaluated seven divenity indices (Shannon-Wiener, Berger Parker, evenness, richness, alpha, Q-statistic and Margelef, see Chapter 2 for calculations), for comparative purposes. RESULTS

Spiders

Doane & Dondale's (1979) study reported that alpha, the Brillouin index and evenness were higher in spider populations within a wheat field than in the field border. Wth aie exception of the Berger-Parker index (0.30 for spiden within the field and O. 15 for spiders at the border), my re-analysis of their data generally supports their conclusions (Table 5-1). Thrir data were re-examined for fit to divenity and abundance models. For both sites, combined data for the two study yean statistically confoned to log series distributions (X2-4.67, df =8, ps0.50 for the within field population and X-2.17, df =8, p0.50 for the border population) (Figure $la). Fit to log nomal distributions could not be tested because then was no mode in the distributions (see Chapter 4 for rationale).

Coyk (1981) examined spider populations in forested and three clearcut areas of varying ages. He used the Shannon-Wener index, richness and evenness, and reported that clear cutting reduced richness yet increased evenness and species diversity. With the exception of the BergeMarker and evenness indices, my re-analysis of his data supports his condusions (sw Table 51). 1 found that when I fit the data to divenity and abundance models, al1 the sites conforniad to log series distributions of species and abundance (Figure 54b). The mature forest (Ellicott Rock forest) had a value of $=4.36, dk7, pr0.20, the site that had been cut 5 yean prior to the study (Horse Cover clearcut) had a value of $ =4.90, df=7, p0.20, the site which had been cut 2 yean prior to the study (Ellicott Rock deanut) had a value of f-10.32, df =6, p0.10, and the site which had been cut 1 year before the study (Buck Creek dearcut) had a value of Tak 51. Oiversity indices for species data in published studies. A is abindance, R is richness, H' is the Shannon-Wiener index, E is wenness, 6-P is the Berpr-Parker index, Al is alpha diveruty 0 is the O-statistic and M is the Margelel index.

omwbuwo TYP. sia -ripli- ~kkgkil~~~ipA R H' E B-P M O AI Rohtonco bspins Old Gr& LeioadBeetles 3489 32 237 0.M 0.30 3.80 17.16 4.87 PedcandCtmdw 1934 ~b9m4o~~sag0 1235 32 2.58 0.74 0.25 4.35 19.52 6.00

Mare toresi (~iiioott~ock) Spiders 595 tio 3.17 0.77 0.16 9.24 41.91 9.76 coyie 1961 2 yean sin- dear cut (Ellicot! Rock) 297 53 3.41 0.86 0.13 9.13 22.W 9.32 1 year since dear cut (Buck CI&) 419 50 3.21 0.82 0.13 8.12 17.15 7.00 5 yeers shce &ar cut (Horse Cove) ' 418 71 3.65 0.86 0Zû 11.60 29.14 10.30

Of Ruts Mites On Rits

Field Bord01

Plawed - annual Spiders 84 18 2.17 0.63 0.01 3.84 13.59 7.03 Haslans and Shaddy 1996 Mowmd-mual 133 12 2.60 0.75 0.03 2.25 15.50 3.20 Bumed - mnuai 213 26 2.58 0.74 0.21 4.66 16.32 7.77 Pkwed - succession 214 35 2.89 0.83 0.21 6.34 26.34 11.89 control 210 31 2.76 0.80 0.22 5.61 20.71 10.04

Old field bes 101 54 128 2.64 0.76 0.43 13.77 30.99 1 1 ,O1 MacKay and Knerer 1979

ChemicaWesticide Fields with exposure to Fenitrothbn Bem 129 28 0.62 0.18 0.15 5.56 25.75 10.33 Kevan et ai. 1997 FMno exposure Io Fenitothion 1174 49 0.62 0.18 0.15 6.79 31.04 15.83

Pm Bt beatment LepldoQteta 831 63 3.23 0.93 0.16 9.22 49.00 10.26 Bu- et al, 1995 Bt treaùnent 163 29 2.85 0.82 0.17 5.50 18.93 11.62 Post Bt beatment 246 36 2.07 0.60 0.52 6.36 35.63 14.11 Fig. 5-1. Rank abundance plots of data sets included in my meta-analysis on a logarithmic scale against the species' rank, in order from the most abundant to the least abundant species.

5.1 a Spider Data from Doane and Dondale (1 979)

0.1 I 1 t 1 21 31 41 Species saqwncs 5.1 b Spider data from Coyle (1 881)

-Ellkot Rock Forest

4ElliaAt Rock Clearcut +Buck Creek Clearcut 3 1 -O- Hom Cowr Clerrcut

0.1 1 11 21 31 41 51 61 71

5.lc Spider data fmm Haskins and Shaddy (1 986) Fig. 5-1 cont. Rank abundance plots of data sets included in my meta-analysis on a logarithmic sale against the species' rank, in order from the most abundant to the least abundant species.

5-ld Diving beetle data fmm Nilsson and Soderberg (1 996)

5.1 e Soil mite data from Kevan et al. (1995)

lmol

+off rut +on track Fig. 5-1 cont Rank abundance plots of data sets included in my meta-analysis on a logarithmic scale against the species' rank, in order from the most abundant to the least abundant species.

5.lg Bee data (rom MacKay and Knerer (1979)

5.lf Lepidoptera data from Butler (1992) for pre-81, and Butler et aL(1995) for treatment years $=2.67, dk5, ~~0.75).Fit to log nonnal distributions could not be tested because there was no mode in the distributions.

Haskins & Shaddy (1988) investigated the effeds of buming, mowing and pkwing on spiders in an old field ewsystem. They used the Shannon-Wener index to test for diversity differences between sites and found that divenity was lowest in the annual plowed fields. W~ththe exception of Berger-Parker and alpha indices, my re-analysis of UMir data with other divenity indices support their conclusions (Table SI). I found that when I fit the data to divesitty and abundance models, al1 the sites confonned to log series distributions (Figure 5-lc). The populations confonn to log series distributions (e.g., in the control site x2=3.99,df =5, p+0.25, in the plowed site x2=5.37,df =SIpr 0.25, in the mowed site x2=2.13, df -5, p>0.50). Fit to log normal distributions could nat be tested because there was no mode in the distributions.

Diving ûeetler

Nilsson & Soderberg (1996) examined species richnesr and abundance of diving beetles in temporary water bodies, and in permanent lakes. My re-analysis of their data wi-th additional divenity indices indicates the divenity of beetles in permanent lakes was higher than in tempomry ones for al1 indices except for the Berger-Parker index (Table 5- 1). I found that when I fit the data to divenity and abundance models al1 the sites confomed to log series distributions (Figure 5-id). In both cases there was confomity to log series model (X-3.62. df =5, p30.50 and $=2.88, df =6, p0.50 reopectively). Fit to log normal distributions could not k tested because them was no mode in the distributions. Tundrr Mites

Kevan et al. (1995) reported abundance, and dchness, of soi1 mites which were wllected on, and off, vehicular ruts in the Arctic. They found that on ruts, abundance and richness were lower Vian off the ruts. My re-analysis of their data with additional diversity indices shows that some divenity indices are higher off the ruts than on the ruts (e.g., richness and Berger-Parker index) whenas other indices are higher on the ruts than off the nits (e.g., Shannon-Wmer index , Q-statistic and alpha (see Table 5-1)). I found that when I fit the data to divenity and abundance models. the mites confomed to a log series distribution under both conditions (x2=7.42, df =8, p>0.25 and $=II.08, df =12, ~~0.25respectively). The distributions are not significantly different from log normal off the rut (x2=8.87,df =12, p>0.13) (Figure 5-le). Fit to log normal distributions on the rut could not be tested because there was no mode in the distributions.

Lepidoptera

Butler et al. (1995) examined the effect of Bacillus thunngiensis (a bacterial produd used for control of gypsy moth lawae) on macrolepidopteran lawae in West Virginia from 1990 to 1992. They found that ahdance and richness of the Lepidoptera were highest in the pre-spray year. With the exception of Berger-Parker and alpha indices, my re-analysis of their data support their conclusions (Table 51). In the yean before and during the treatment the populations confonned to log series distributions p=6.82, df =8, ps0.25 and 2-5.14, df = 5, p+0.25 respectively). In the year following treatment Me distribution did not confonn to log series ($=14.45, df = 5, ~~0.05)but did msemble a geometric distribution (Figure 5-19 becwse of the high nurnber of rare species which were present. It is possible to speculate that the increase in rare species

89 represents a rebound in species nurnber. Log nonnality could not be tested for al1 three periods (befon, durkig and after) because them was no mode in species distributions. Butler (1992) conducted a basrline study of mawlepidoptenn populations during 1984 and 1985 before the gypsy moth infestation moved into West Virginia. Those populations confomed to log series distributions of species and abundance (x2=4.20,df =8, ps0.25) (Figure 5-If). Log normality could not be tested because there was no mode in spedes distributions.

MacKay & Knerer (1979) examined wild bees in an old field habitat in southem Ontario. Their data confomed to log normal distributions of species and abundance ($=O. 17, df =l3, p=0.20) but not log series distributions ($=217.04, df=l3, p

DISCUSSION

Divenity Measures

Re-analysis of the data sets with a wide anay of diversity measures generally supports the ides that higher divemity is present in sites with less disturbance. There is inconsistency between different indices, as was diswssed in Chapter 2, with respect to both my carabid study and meta-analysis. This supports my conclusions in Chapter 2 that divenity indices do not necessarily provide reliable tools for detecting disturbance. For carabid studies presented in Chapter 2, as well as the studies in this chapter, species richnesr and abundance tend to be higher in undisturbed sites. May (1975) suggests that spscies richnesi is the kstmeasum of a spedes assemblage. My studies do not 90 dispute May's suggestion, however, richness alone is not useful because richness perse provides little information on ewsystem fundon. As well, with richness, inter-site and inter-time comparisons are needed to detemine the effect of disturbance, if any. The advantage of deviation from log nonal distributions of divenity and abundance, as discussed in Chapter 1, is that there is a theoretical standard against which to make comparisons, and inter-site and inter-time comparisons are not needed.

Spiden and Diving htks

Common features between these invertebrate groups are that they are entirely predaceous and that they confom to log series distributions. However, both spiders and diving beetles can be associated with other invertebrate species that have overlapping feeding niches with them (e.g., for diving beetles, the Noteridae (Bonor et al. 1989). Thur, it is likely that spiders and diving beetles are not single functional groups.

For spiders Me situation is more complex than with diving beetles because there are different hunting strategies within spider species assemblages (e.g., web builders, ground hunten etc.). Thus, the divenity and abundance analysis done in this chapter may not be appropriate because: 1) the data sets used in this chapter grouped al1 spiders together, possibly combining several different functional groups, and 2) there are not enough spider species reprewnted within a feeding niche to test how disturbance might affect the functional group.

Soil Microrrthmpods

Tundra soi1 mites appear to be good indicators of disturbance when deviation

91 from a log normal distribution of species and abundance is used as an indicator. However, thb conclusion must be accepted with caution because log normal distributions of the "on rut" mite assemblage could not be assessed because there was no mode in species distributions. Accepting tundra mites as a good gmup for disturbance studies in the arcüc would bo easy because vehide track disturbance is obviously very disniptive to arcüc ecosystems (Kevan et al. 1995). One of the pmblems with mites is that there is more than one fundional group within the mite assemblage (e.g., detritivores, herbivores, predaton) (Borror et a/. 1989). Fumer study is required before concluding that mites are useful indicators of disturbance.

HBgvar (1994) reported that soil microarthropod divenity and abundance structures were sensitive to disturbances such as acidifiaon and heavy metals. He examined both Collembola and mite assemblages and found they deviated from log normality under conditions of disturbance. Oominance graphs provided in Peterson's (1995) study suggests that collembotan assemblages deviate from log normality when exposed to agriculturcil cultivation. Collembola are primarily detritivores (Bonor et al. 1989)

Taxomomic, bionomic or numerical constraints did not limit soi1 mite and collembolan assemblages in these studies. These assemblages meet some, but not all, the fundional constraints that were detenined to be necessary from by my carabid study (see Chapter 4). The conditions that need to be met are that feeding behaviour is not size dependant (for CoHembola, but not necessarily for soi1 mites), as well, that thmughout their Iife stages the arthropod species should have similar behavioun. As with carabids, spiders and diving beetles, soi1 mites are a collection of partial fundional groups. Collembola are not the only detritivores present in the soil, thus are a component of the soi1 detritivore cornplex. Although they are not a complete detritivore taocene, they dominate soi1 ewsystems sufficiently for there to be a mode in the divenity and abundance data. This group may be witable indicaton for disturbance. As with microlepidoptera, bees are good indicaton for disturbance. Kevan et al. (1997.) th& that pollinating Mes deviated from log nomal divenity and abundance patterns when they wcm, exposed to pesticide spraying. However, as A# Lepidoptera populations, some bee populations that am under chmnic human disturbance can ôe desuibd by log normal distributions (MacKay & Knenr 1979).

Neither beets, nor rnacrolepidoptera, are limited by numeric, bionornic, taxonomic (at kast krthese studies) constrakits (see Chapter 1). 00th bees and Lepidoptera have feeding behavioun that are not mtricted by their size. For example, many pollinators, regardless of size, compete for the same resources, and have overiapping niche requirements (Kevan & Baker 1983). Carabid beeUe size, on the other hand, does detemine the size of prey which can be hunted (Loreau 1988). FurVierrnore, generally neithw bees nor adult mscrolepidoptenhed on a wide variety of food types. For example, adult bees are mosUy limited to feeding on nectar and pollen, and most adult macrolepidoptera lawae hed on nectar. This behaviour contrasts with carabid beetles which feed on a variety of food items (see Larochelle 1990 and Appendix 1). In general, bees, as either larvae or adults have similar behaviours within life stages. The same holds tnie for Lepidoptera. How~vW~because adult Lepidoptera have greater feeding niche overlap than do larvae. for considerations of deviation from log normal distributions of diversity and abundance, it may be more appropriate to use adults as indicators of disturbance.

These assemblages reveal the very important point that sometimss a habitat that seems to be distuibed, it is in fad not distuibed to a given functional group. Effects of some distuturbances cannot be measured in ternis of deviation from log normal distributions of divenity and abundance because the indicator groups are already disturbed by an undetertnined force. Leiodid Beetks, Coctimlids and Gnrshopprn

Leiodid beetles confomed to both log series and log normal models (see Chandler & Peck 1992). This finding suggests Mat more work is required on this group. Leiodids are fungivoms, however they may not be a complete functional group on their own. As well, the logging disturbanœ Chandkr & Peck (1992) studied might not have been disniptive to the beeties because the study area had been logged more than four decades prior to the study. Nummelin (1998) explored deviation from log nomality as an indicetor of disturbance by using coccinelids and grasshoppen. He kund that these species assemblages cwld not be tested because they did not have modes (see Chapter 4) and were characterized by log series distributions. These taxa do not comprise complete functional groups.

Carabids, spiders and diving beeUes comprise partial functional groups, and thus are not usehil as disturbance indicaton on their own. They need to be subdivided, and the wmponents pooled with species that have similar functions and constraints within the ecosystem. Leiodid beetles, cooci'nelids and grasshopprrs am parts of functional groups and need to be completed as functional groups before becoming useful disturbance indicaton. In contrast, pollinating bees and adult macmlepidoptera are more or less cornplete functional groups and should be explored as indicaton for disturbance.

Kevan et a/. (19Q5),and Hggvar (1995) both collected mite species in extreme habitats. Kavan et el.'$ (1QQS) sites wem in the high Anüc, and HBgvar (1895) collected his mites from sites mat are highly acidified or polluted with heavy metals. Although the mite collections rnay not represent a single functional group (see above) it is possible that their collections incîuded most species from each mbfunctional gmup. If al1 species, and al1 functional groups. in an ecosystem are pooled together under undisturbed conditions there should br log nomal distribution of divenity and abundance of species (Preston 1980; Sugihara 1980). Likewise, if al1 species, and al1 fundional groups, in an ecosystem are pooled together, when an anthmpogenic disturbance occun there would be deviation from log nomality.

It is interesting that two groups, pollinaton and detritivores, which appear to make good disturbance indicators have keystone roles in the ecosystem (Kevan 1991). This suggests that the groups that should be tergeted for disturbance studies are keystone groups which include pollinaton, and detritivores. Indicritor groups may not reflect a change in divenity and abundance distribution if the disturbance that is being tested does not affect assemblage structure either because the disturbance has been in place long enough for the assemblage to be adapted to it (e.g., MacKay & Knerets (1979) bees), or bewuse the assemblage is already disturbed (e.g., Butler et al.'s (1995) Lepidoptera). Deviation of functional gmups from log normal distributions of divenity and abundance is a tool for describing disturbance but it rnust be used with caution. When deviation occun, it raises a red flag that indicates that the fundional group in question may have been disnipted. CHAPTER 6: SYNTHESIS, LIMITATIONS AND FUTURE OlRECTlQNS FOR ASSESSINO DISTURBANCE

In order to study disturbance, ecologists often need to measure species diversity. I chose to use a gradient of tillage, and 14 different rneasures of divenity, to assess the effect of disturbance on carabid beetles. I used two different approaches: divenity indiœs (Chapter 2), and deviation from divenity and abundance models (e.g., species size divenity (Chapter 3) and species divenity and abundance (Chapter 4)). 1 found that none of the seven divrnity indices I used (richness, Shannon- Weiner, Berger-Parker, Q- statistic, Margalef, alpha and evenness) indicated that carabid beetles were affected by tillage disturbance. Although this supports the findings of some authors (Tyler 8 Ellis 1979; Dritschillo & Ewn 1982; Hokkanen & Holopainen 1986; Kromp 1989; Mack & Buckrnan 1890; Tonhasca 1993), it wns contrary to the cornmonly held view and findings of othem (Dritschillo & Wanner 1980; House & All1981; House & Stinner 1983; Ferguson

& McPhenon 1985; Brust el el. 1986; Weiss et 81. 1990; CBrcemo 1995). The difference between my study and the mers is that I examined a wide range of divenity indices rather than depending on the outcome of one or two. This is the first tirne that such a large amy of indices has been used as cornparisons for a single data base.

My conclusions that cerabid beetles are not affeded by tillage is also supported by diversity and abundance models. In Chapter 2, 1 Lnvestigated the effect of tillege on cambid size as a measure of ecological fundion, and did not find significant differences among treatments (i.e., fenns which represent different degrees of tillage). This holds true for al1 four measums of body size which I used (man Iength, divenity of sizes, distribution of length and distribution of biovolume). Although the log normal distributions

97 of length, and biovolume suggest that carabid beetles are a functional group, it b possible that these results anmerely statistical artifads. An alternative explanation is that carabid sizes ais log nomally distributed kcause size distributions represent niche hierarchy. Body riz@might represent functional pmperües of species that accurately refiect niche hierarchy. This idea needs fumer investigation, but details of species ecology and niche requirements must fint be detemined to establish if, in fact, size does reflect functional properties. Because Viis is a major undertaking, and other species assemblages offer greater promise as disturbance indicaton (see Chapter 5), this idea should not be a research priority. Nevertheless, because carabid size distributions did not deviate hmlog nomal distributions with increasing disturbance, this approach may not be useful for assessing disturbance.

I was not able to conclude that carabid beetles are a functional group by using species diversity and abundance distributions of species. Even in undisturbed conditions carabid beetle assemblages are not distributed log nonnally. This indicates that niche hierarchy does not occur within carabid beetle assemblages, and thus, that carabid beetles are not a functional group. It is likely that then is little intenpecific competition among carabids because they have diverse feeding behavioun (Thiele 1977; Larochelle 1990) and size dependant feeding (Loreau 1988; Rushton d al. 1991). Carabid beetles probably ought to be divided into size restricted subgroups, and then combined with spcies from other taxa which sham similai sizes and feeding behavioun. This suggestion is a major undertaking because the ecological inter-relationships of most invertebrate species am not well nsearched. Befom this suggestion can be acted upon, the ecology of most of the potenüal carabid, and non-carabid species within a sub-group, would have to be detemined. As well, because carabid divenity and abundance patterns are not log normal, even undri undisturbed conditions, they are not useful for assessing disturbance when deviation away from log normality is wnsidered. The above condusions from my study of carabid beetles. which are drawn from divenity indices, and diversity abundance models, are supported by meta-analyses of forty five published data sets (SM Chapter 4). 1 did not find any one divenity index to be mon useM than another in pin-pointing disturbance. As well, carabid beetle assemblages wen not useful when using deviation fmm log norrnality as an indication of disturôance. Canbid beetle assemblages are log series, or geornetrically, distributed in both disturbed and undisturbed sites.

Meta-analyses of nonarabid species assemblages indicate that groups which should be targeted for disturbance studies are keystone groups which indude pollinators, and detritivores (se8 Chapter 5). Although I conclude that deviation of functional groups from log normal distributions of diversity and abundance is a useful tool for describing disturbance, it must be used with caution. When deviation occurs, it raises a red flag that indicates that the functional gmup in question rnay have been disnipted. FurVler, if the disturbance that is being tested is found not to affect community structure, that may result because the disturbance has been in place long enough for the cornmunity to be adapted to it, or because the cornmunity is already somehow disturbed.

LIMITATIONS Them are two main limitations to my study:

1) My ewpefimental design is limited by Vis number of fams I used. It would have been preferable to have used several farms in each of several different geognphic locations for making inter-comparisons between fams as repmsentative of tillage. If that had been possible, from r practical and cost stand point, then stronger inferences about the rok of tillage disturôance on carabid beeties wuld have been made. 2) Although deviation from log normal distributions of diversity and abundance has been shown to be useful for other specires assemblages, it was not useful for carabid beetîes, in dther analyses of rny own nsults or Hi meta-analysis of the results of others. fhat I had to aaccpt my nuIl hypotheses is an interesting and valuable finding. However, when nuIl hypotheses cannot be rejected, it is often more difficult for the work to be accepted.

FUTURE DIRECTIONS

If carabid beetles are to be used in assessing effects of disturbance, their ecology together with that of other invertebiates, needs to be elucidated to detemine which species can be grouped within the appropriate carabid sub-group in order to fonn a complete fundional group. Hypotheses regarding sire distributions of carabid beetles, and whether or not their sizes reflect niche hierarchy, should also be further explored. As ml, species assemblages other than carabid beetles. for example bee pollinaton and Lepidopteta, could be used to further study the hypoaiesis that with increasing disturbance there are changes in body sizes within the assemblage.

A more theoretical direction which should be taken is to determine the time it takes from an initial disturbance befors a fundonal group returns to e log normal distribution of divenity and abundance. Given that the effects of disturbance on ecosystems must be measured by changes in aie processes in the systems, direct, or indirect mersures, of the processes rnust be made. Cornparisons with divenity indices between disturbed and undisturbed systems, and gradients of disturbance are valuable, but require spatial or temporal wmparisons. One way to ovetcome the shottcomings of divemity indices is to have a divenity measura which is ustanbalone"and does not need spatial or temporal comparisons. With such a stand-alone measure, a theoretical standard against which to maka compsrisons is mquind. I used log normal distributions of divenity and abundana as a stand-alone measure because the distribution is based on cornpetitive exdusion and niche hierarchy. One of the future direcü*ons of my research is to detemine if then are stand-alone measuns othw than log nomal distributions of diversity and abundanw. All potential measures must be besed on ecological and evolutionary pnnciples, such as cornpetitive exclusion, niche hierarchy, resilience (Krebs 1985), or succession (Krebs 1985). This approach to measuring the effect of disturbance is in its infancy.

My work has attempted to use indices and models on species, and species assemblages. My results indicate neither approach works reliably on carabids. Nevertheless, my meta-analysis of non-carabid assemblages indicates that there are gmups, such as pollinating bees and macrdepidoptera, which show promise as good indicators of disturbance. Further research is needed to find other useful indicator gmups. REFERENCES

Allen, R.T. (1976) The occumnce and importance of ground beetles in agricultural and sunounding habitats. In Carabid Beetles: Their Evolution, Naturel History and Clessiifcation (eds T.L.Emin, G. E. Balla D.R. Whitehead) pp. 485-505. Pfoceedings of the First International Symposium of Carabidology. Smithsonian Institution, Washington, D.C. Allmaras, R.R. & Dowdy, R.H. (1985) Conservation tillage systems and their adoption in the United States. Soi1 Tillage Research, 5, 197-222.

Andrewartha, HG. & Birch, L.C. (1954) The Distdbution and Abundance of Animals. University of Chicago Press, Chicago Arthur, W. (1984) Mechanisms of Mo~hologicalEvolution: A Combined Genetic, Developmental and Ecological Appmach. John Wiley and Sons, Chichester. Asteraki. E.J.. Hanks, C.B. 8 Clements, R.O. (1992) The impact of the chernical removal of the hedge-base Rora on the comrnunity structure of carabid beetles (Col., Carabidae) and spiders (Araneae) of the field and hedge bottom. Journal of

Appried Entomology, 1 1 3, 396406.

Baines, D., Stewart, R. & Boivin, G. (1990) Consumption of carrot weevil (Coleoptera: Curculionidae) by five species of carabids (Coleoptera: Carabidae) abundant in carot fields in southwestern Quebec.Envimnmental Entomology, 19, 1 146-1 149.

Bamey, R.J. 4% Pass, B.C. (1988a) Gmund beetle (Coleoptera: Carabidae) populations in Kentucky alfalfa and influence of Mage. Journal of Economic Entomology, 79, 511-517. Bamey, R.J. & Pass, 8.C. (1986b) Foraging behaviour and feeding preference of ground beetîes (Coleoptera: Carabidae) in Kentucky alfalfa. Journal of Economic Entomo/ogy, 79, 1334-1 337. Belaoussoff, S. 8 Kevan, P.G. (1998) Toward an ecological approach for the assessrnent of ecosystem health. Ecosystem Health, 4,408. Best, R.L. 8 Beegle, C .C. (1977a) Food pnferences of five carabids comrnonly found in

lowa comf ields. Environmental Entomology, 6, 9-1 2.

Best, R.L. 8 Beegle, C.C. (1977b) Consumption of Agmstis ipsilon by several species of carabids found in lowa. Environmental Entomology, 6. 532-534.

Borror, D.J.. Triplehom, C.A. 8 Johnson, N.F. (1989) An Introduction to the Study of lnsects, 6'"dition. Saunden College Publishing, Orlando, Çlorida. Boswell, M.T. 8 Patil G.P. (1971) Chance mechanisms generating the logarithmic series distribution used in the analysis of numbers and individuals. In Statistical Ecoiogy

(eds G.P. Patil, E.C. Pielou & W.€. Waters) pp. 99-130 Pennsylvania State University Press, University Park, PA. Bousquet, Y., Smetana, A., 81 Madison D.R. (1984) Trechus quadristriatus, a palabarctic species introduced into North America (Coleoptera: Carabidae). The Canadian Entomologist, 11 6, 2 15-220. Boyd, H.P. (1978) The tiger beetles (Coleoptera: Cicindelidae) of New Jeney with special reference to their ecological relationships. Trensactions of Ihe American

Entomological Society, 104, 19 1-242.

Brown, D.M. 8 Bootsma, A. (1993) Crop Heaf Units for Corn and other Wann-season Crops in Ontafio. Ministry of Agriculture and Food Publication 93-1 19.

Brown, J.H. (L Munger, J.C. (198s) Experîmental manipulation of a desert rodent community; food addition and species removal. EcoIogy, 66, l54W563.

Brown, J.H. 8 Maurer. B.A. (1986) Body size, ecology dominance and Cope's rule. Nature, 324,24&250.

Bnist, G.E. 8 House, G.J. (1988) Weed seed destruction by arttiropods and mdents in low-input soybean agroecosystems. AmrFcen Journal of Alternative Ag~cultum, 3,19-25. Bwst, CE.. Stinner, BR. 8 McCartney, D.A. (1986a) Predation by soi1 inhabiting arthropods in intemopped and monoculture agroecosystems. Agncultum, Ecosystems and Environment, 18, 145-1 54.

Brust, GE.,Stinner, BR. & McCartney. D.A. (1986b) Predator activity and prudation in corn agroecosystems. Envimnmental Entomology, 15, 101 7-1 02 1. Butler, L. (1992) The community of rnacrolepidopterous larvae at Coopers Rock State

Forest. West Virginia: A baseline study. Canadien Entomologist, 124, 1149: 1156. Butler, L., Zivkovich C., 8 Sample RE. (1995) Richness and abundance of arthropods in the oak canopy of West Virginia's eastern ridge and valley section during a study of impact of Bacillus thuringiensis with emphasis on macrolepidoptera lanrae. Reporl to West Viqinia Uniwtsity, Agricultural and Fomst Experiment Station.

Carcarno, H.A. (1995) Effect of Mage on ground beetles (Coleoptera: Carabidae): A fan-scale study in central Alberta. The Canadian Entomologist, 127, 631-639. Carcarno, H.A.. Niemela, J.K. & Spence, J.R. (1995) Faming and ground beetles: effects

of agronomie practice on populations and community structure. The Canadian Entomologist, 127,123-1 40. Carpenter. S.R. (1990) Large sale peflurbations: Opportunities for innovation. Eco/ogy, 71,2038-2043.

Chadrasekani. W.U. 8 Frid, C.L.J. (1996) Effect of human trampling on tidal Rat fauna. Aquatic Consetvation: Marine And Fmshwater Ecosystems, 6,299-331.

Chandler, O.S. & Peck, S.B. (1992) Divenity and seasonality of leiodid beetles (Coleoptera: Leiodidw) in an old-growth and a 40-yearsld forest in New Hampshire. EnMmnmentel Entomdogy, 21,f 283-1293. Clark, f. & Enter, D. (1984) Vegetation disturbance caused by cottage disniption in central Ontario. Journal of Envimnmenttel Menagemnt, 18. 229-239. Clark, M.S., Luna, J.M., Stone, N.D. & Youngman, R.R. 1994. Generalist predator . consumption of amywon (Lepidoptera: Noctuidae) and effect of predator removal on damage in no-till corn. EnvironmentalEntomology, 23, 817-622. Colinvaux . P.A. (1973) Introduction to Ecology, John Wiley 8 Sons. New York. Colwell. R.K. & Winkler, D.W. (1984) A nuIl model for nuIl models in biogeography. Ecological communities: Conceptual issues and the evidence (eds O.R . Strong, O.,Simberloff, L.G. Abele & A.B. Thisle,) Princeton University Press, Princeton, New Jersey. Comell, H., Hurd, L.E. & Lotfich, V.A. (1976) A measure of response to perturbation used to assess structural change in some polluted and unpolluted stream fish communities. Oecdogia, 23, 335-342. Coyle, F.A. (1981) Effects of clearcutting on the spider community of a southem Appalachian forest. Journal of Anchnology, 9, 285-298.

Crain, 1. K. (197 1) Possible direct causal relation between geomagnetic reversais and biological extinctions. Bulletin of the Geological Society of America, 82, 2602- 2606. Critchley, B.R., (1972) Field investigations on the effects of an organophosphonis pesticide, thionazin. on predacious Carabidae (Coleoptera). Bulletin of EntomologÉcal Research, 62, 327-342.

Davies, M.J. (1953) The contents of the crops of some British Carabid beetles. The Entomobgist3 MontMy Magazine, 89,18-23. Davies, M. (1959) Contribution to the ecology of species of Notiophilus and altied genera

(Col., Carabidae). The Entomologist's Monthly Magazine, 91, 25-28. Davy, A. J., Noble, S.M. & Oliver, R.P. (1990) Genetic variation and adaptation to flooding in plants. Aquetic Botany, 38, 91-108. Davey, R.H., Murphy, S.M. Wiens J.A., Hayward, G.D., Hamer, E.J. 6 Smith, L.N. (1997) Effects of the Exxon Valdex oil spill on habitat use by birds in Prince William

Sound, Alaska. Ecological Applications, 7, 593-6 13. den Boer, P.J. (1980) Exclusion or coexistence and the taxonornic or ecological

relationship between species. Netherlends Journal of Zoology, 30 , 278-306.

Desender. K. 8 Pollet, M. (1985) Ecological data on Clivina fossor (Coleoptera, Carabidae) from a Pasture ecosystem. II. Reproduction, biometry, biomass, wing polymorphism and feeding ecology. Revue d'Ecologie et de Biologie du Sol, 22,

233-245. Dewdney, A.K. (1997) A dynamical model of abundances in natural communities. Coenoses, 12,67-76.

Dillon, E.E. 8 Dillon, L.S. (1961) A Manual of Cornmon Beetles of Eastern Noflh America. Evanston, Illinois Peterson.

Digweed, S.C., Cume, C.R., Carcarno, H.A. & Spence, J.R. (1995) Digging out the "digging -in effect" of pitfall traps: Influences of depletion and disturbance on catches of ground beetles (Coleopteta: Carabidae). Pedobio/ogia, 39, 561-576.

Doane, J.F. & Dondale, C.D. (1979) Seasonal captures of spiders (Araneae) in a wheat field and its grassy borden in central Saskatchewan. The Canadian Entomo/ogist,

il 1, 439-445. Dritschilo, W. 8 Erwin, T.L. (1982) Responses in abundance and divenity of cornfield carabid communities to differences in farrn practices. Ecology, 63, 900-904.

Dritschilo, W. & Wanner, D. (1980) Ground beetle abundance in organic and conventional corn fields. Environmental Enfom~lugy~9,629431. Dufrdne, M. & Legendre, P. (1997) Species assemblages and indicator species: f ha nwd for a flexible asymrnetrical approach. Ecological Monogrephs, 67, 345-366. Edwards, C.A., Sunderland K.O. & George K.S. (1979) Studies on polyphagous predaton of cereal aphids. Journal of Applid Ecology, 16,811-823. Ehrenfeld, D.W. (1972) Consewing Lih on Earlh. Oxford University Press, New York. Elton, C. (1958) The Ecology of Invasions by Animais end Plants. Methuen, London.

Erwin,T.L. , G.E. Bal! & Whitehead D.R.(eds) (1976) Carabid Beetles: Their Evolution, Natuml History and Classification. Proceedings of the Fint International Symposium of Carabidology. Smithsonian Institution, Washington, D.C.

Eyre, M.D., & Ruston, S.P. (1989) Quantification of conservation criteria using invertebrates. Journal of Applied Ecology, 26, 159-171.

Fan, Y., Liebman. M. Groden, E. & Alford, A.R. (1993) Abundance of carabid beetles and other ground-dwelling arthropods in conventional versus low-input bean cropping systems. Agricultum, Ecosystems and Environment, 43, 12% 139.

Ferguson., H.J., & McPhenon, R.M. (1985) Abundance and divenity of adult Carabidae in four soybean cropping systems in Virginia. Journal of Entomological Science, 20,163-171.

Fisher, R.A.. Corbet A.S. & Williams C.B. (1943) The relation between the number of species and the number of individuals in a random sample of an animal population. Journal of Animal Ecology, 12, 42-58. Floate, K.O., Doane, J.F. & Gillott, C. (1990) Carabid predaton of the wheat midge (Oiptera: Cecidomyiidae) in Saskatchewan. EnMmnmtal Enbmology, 19.1 505- 1511. Forsythe, TG. (1983) Mouthparts and feeding of certain ground beetles (Coleoptera: Carabidae). Zoologka/ Journal of the Linman Society, 70, 31 9-376. Frank, J .H . (197 1a) Carabidae (Coleoptera) as predators of the red-backed cutwom (Lepidoptera: Noctuidae) in central Alberta. The Canedian Entomologist, lO3,lO39-lO44.

Frank, J.H. (1971 b) Carabidae (Coleoptera) of an arable field in central Alberta. Quaestiones entomologicae, 7,237-252. Fraser, J. (1980) Acclimation to lead in the freshwater isopod Asellus aquaticus.

Oecologica, 45, 4 19-420.

Freitag, R., Hastings, L.. Mener, W. R. & Smith, A. (1973) Ground beetle populations near a kraft mill. The Canadien Entomologist, 105, 299-310. Fretwell, S. (1978) Cornpetition for discrete venus continuous resources: tests for predictions from MacArthur-Levies models. American Natumlist, 1 12, 73-81. Gaver, U. (1997) Collembola in Central Amazon innundation forests - Strategies for suwiving floods. Pedobiologia, 41,69-73.

Gee, J .H.R. & Giller P.S. (1986) O~anizationof Communities: Past and Pliesent. BlackweH Scientific, Oxford. Glen, W. (1990) What killed the dinosaun? Ameiican Scientist, 78, 354-370.

Graves, R.C. (1983) The Cicindelidae of Michigan (Coleoptera). American Miûland

Naturalist, 69, 492-507.

Graves, R.C. (1965) The distribution of tiger beetles in Ontario (Coleoptera: Cicindelidae). Pmcwdings of the Entomo/ogica/ Society of Ontaiio, 95,63-70.

Gray, J.S. (1983) Use and misuse of the log-nonal plotting method for detection of effects of pollution - a reply to Shaw et al. (1983). MaRne Ecdogy - Pmgmss Senes, 1 1 , 203-204.

Gray, J.S. (1979) Pollution-induced changes in populations. Philosophica/ Transactions

of the Royal Society of London. B., 286,545-561.

Gray, J.S. & Mirza, F. B. (1979) A possible rnethod for the detection of pollution-induced disturbance on marine benthic wmmunities. Marine Pollution Bulletin, 10,142-

146.

Griffiths, D. (1988) Size-abundance relations in communities. Amaan Nalumlist~127,

140-1 66. Hagley, E.A.C., Holliday, N.J. & Barber, D.R. (1982) Laboratory studies of the food preferences of some orchard carabids (Coleoptera: Carabidae). The Canadian

Entomologist, 114,431 -437. Hdgvar, S. (1994) Log-normal distribution of dominance as an indicetor of stressed soi1 micfoarthropod communities? Acta Zoologica Fennica, 195, 71-80

Halme & Niemela (1993) Carabid beettes in fragments of coniferous forest. Annals

Zoologica Fennica, 30, 17-30. Hairston, N.G. 8 Byers, G.W. (1954) The soi1 arthropods of a field in southem Michigan: a study in community ecoiogy: Contributions of the Labomtory for Veflebrate

Biology of the University of Michigan, 64.1-37. Hance, T. 8 Gregoire-Wibo, C. (1987) Effect of agricultural practices on carabid beetle populations. Acta Phytopathologica Entomologica Hungary, 22, 147-7 60.

Harvey, P.H. & Godfray, H.C.J. (1987) How species divide resources. Amencan

Naturalist, 129, 3 18-320.

Harvey, P.H. 8 Lawton, J.H. (1986) Patterns in three dimensions. Nature, 324, 212.

Haskins, M.F. & Shaddy, J.H. (1986) The ecological effects of buming, mowing, and plowing on ground-inhabiting spiders (Araneae) in an old-field ecosystem. The Journal of Atachnology, 14, 1-14.

Hejkal, J. (1985) The development of a carabid fauna (Coleoptera, Canbidae) on spoil banks under conditions of primary succession. Acta Entomologica

Bohemoslovaca, 82, 321-346.

Hengeveld, R. (1980) Qualitative and quantitative aspects of the food of ground beeties (Coleoptera, Carabidae): A review. Nethedands Journal ofZoo/ogy, 90, 555-563. Hill. J.M. & Knisley, C.B. (1992) Frugivory in the tiger beetle, Cichdela mpanda (Coleoptera: Cicindelidae). Coleopterists Bulletin, 46, 3-3 10. Hill. J.K., Hamer, K.C., Lace, LA. & Banham, W.M.T. (1995) Effects of selactive logging on tropical forest buttemies on Buru, Indonesia. Journal of Applied Ecobgy, 32, 754-780. Hill, J.K. & Hamer, K.C. (1988) Using species abundance rnodels as indicators of habitat

disturbance in tropical forests. Joumsl of Applied Eco/ogy, 35,458-400. Hobbs, R.J. & Huenneke, L.F. (1992) Disturbance, diversity and invasion: Implications for conservation. Conservation Biology, 6, 324-337.

Hokkanen, H. & Holopainen, J.K. (1986) Carabid species and activity densities in biologically and conventionally managed cabbage fields. Journal of Applied

E~I~o~o~o~Y,102, 353-363.

Holliday, N.J. & Hagley E.A.C. (1978) Occurrence and activity of ground beetles (Coleoptera: Carabidae) in a pest management apple orchard. The Canadian

Enlomologist, 11 O, 11 3- 11 9.

House, G. J. & Ail, J.N. (1981) Carabid beetles in soybean agroecosystems. Environmental Entomology, 10,194-196.

House, G.J. & Pamelee, R.W. (1985) Companson of soi1 arthropods and earthwoms

from conventional and no-tillage agroecosysterns. Soi1 and TiIIage Research, 5, 351-360. House, G.J. & Stinner, B.R. (1983) Arthropods in no-Mage soybean agroecosystems: Community composition and ecosystem interactions. Environmental Management, 7,23-28. Hutchinson, G.E. (1953) The concept of pattern in ecology. PTocBBdiings of the National Academy of Science, PhiIadeIphia, 1 OS, 1- 12. Hutchinson, GE. (1978) An lntmcfuction to Population Ecology. Yale University Press, New Have, Connecticut. Izsak, J. & Hunter, PR. (1992) Serotype abundance distributions in reports of Sa/mone/la incidents in domestic livestock as indicaton of the population biology of

SalmonelIa infections. Functional Ecology, 6, 154- 159. ~aksi6,F.M. (1981) Abuse and misuse of the bn "guild" in ecological studies. OIKOS. 37, 397-400. Jarman, P.J. (1974) The social organization of antelope in relation to their ecology. Behaviour, 48,215-267. JaroSik, V. (1983) A companson of the diversity of carabid beetles (Col., Carabidae) of two floodplain forests differently affected by emissions. V&t 6s Spole6Zoologica. 47, 2 15-220. Johnson, N.E. & Cameron, R.S. (1969) Phytophagus ground beetles. Annals of the

Entomological Society of Amrice, 62, 909-914.

Judd, K.W. & Mason, C.F. (1995) Colonkation of a restored landfill site by invertebrates, with particular reference to the Coleoptera. Peaobiologia, 39, 116-125. Kaulbars, M.M. 8 Freitag, R. (1993) Geographical variation, classification, reconstnicted phylogeny , and geographical history of the Cicindela sexguttata group

(Coleoptera: Cicindelidae). The Canadian Entomologist, 125, 267-3 16.

Kempton, R.A. 8 Taylor, L.R. (1974) Log-senes and log-normal parameten as diversity discriminants for the Lepidoptera. Joumal of Animal Ecology, 43, 381-399. Kerr, SR.(1 979) Prey availability, metaphoetesis, and the size structure of lake trout stocks. Joumal of Fish Research, 43, 187-198. Kevan, P.G. (1991) Pollination: Keystone process in sustainable global productivity. Acta

Hotticu/tume 288,103- 11 O.

Kevan, P.G. and Baker, HG. (1983) lnsects as fiower visiton and pollinaton. Annuel Re&w of EntomoIogy, 28,407453.

Kevan, P.G., Forbes, B.C.. Kevan, S.M. & Behan-Pelletier, V. (1995) Vehicle tracks on high arctic tundra: their effects on the soil, vegetation, and soi1 arthropods. Journal of Applied EcoIogy 32,655467. Kevan, P.G.. Greco. C.F. & Belaoussoff, S. (1997) Biodivenity and abundance in diagnosis and measuring of ecosystern health: pesticide stress on pollinaton on blueberry heaths. Journal of Applied Ecology, 34,1 122-1 136. King, C.E. (1964) Relative abundance of species and MacArthur's model. Ecology, 45, 716-727. Kirchner, TB., Anderson, R.V. 8 lngham R.E. (1980) Natural selection and the distribution of nemotode sizes. Ecology, 61, 232-237. Kirk, V.M. (1971) Biological studies of a ground beetle, Reroslichus lucublandus. Annals

of the Entomological Society of Ametka, 64. 540-544.

Kirk, V.M. (1972) Seed-caching by lawae of two ground beetles, Harpalus pensylvanicus

and H. enaticus. Annals of the Entomo/ogical Society of America, 65, 1426-1428.

Kirk, V.M. (1973) Biology of a ground beetle, Harpalus pensylvanicus. Annals of the

Entornological Society of America, 66, 5 13-5 18. Kirk, V.M. (1975a) Biology of Pterostichus chalcites, a ground beetle of cropland. Annals

of the Entomological Society of America, 68, 855-858.

Kirk, V.M. (1975b) Biology of Stenolophus (=Agonodenrs)comma, a ground beetle of cropland. Annals of the Entomological Society of America, 68,135- 138.

Kitahara, M., & Fujii, K. (1994) Biodiversity and community structure of temperate butterfly species wittiin a gradient of human disturbance: an analysis based on the concept of generalist vs. specialist strategies. Researches on Population

ECOIO~Y,36,187-1 99.

Krebs, C. J. (1985) Ecology: The expen'mental analysis of distribution and abundance. Harper and Row, New York Kromp, B. (1989) Carabid beetle communities (Carabidae, Colaoptera) in biologically and conventionally farmed agroecosystems. AgticuItum, Ecosystems and EnvirPnment, 27,241-25 1. Lambshead, P.J.D., Platt, H.M. & Shaw, K.M. (1983) The detection of differences among assemblages of marine benthic species based on an assessrnent of dominance and diversity. Joumal of Natural History, 17, 859-874.

Laroca, S., Becker, V.O. & Zanella, F.C.V. (1989) Divenity , relative abundance and phenology in Sphingidae (Lepidoptera) in Serra do Mar (Quatro Banos, Pr), south Brazil. Acta Bio/ogica par Cutitiba, 18, 13-53. Larochelle, A. (1990) The Food of Carabid Beetles (Coleoptera: Carabidae, including Cicindelinae). Fabreries, Supplément 5, Association des Entomologistes Amateurs du Québec, Sillery. Québec Lanon, CRS. & MacDonald, G.M. (1998) Fire and vegetational dynarnics in a Jack Pine and Black Spruce forest reconstructed using fossil pollen and charcoal. Joumal of

ECOI~Y~86 8 15,828. Lenski, R.E. (1982) The impact of forest cutting on Me diversity of ground beetles (Coleoptera: Carabidae) in the southem Appalachians. Ecological Entomology. 7, 385-390. Lincoln. R.J., Boxshall, GA. 8 Clark, P.F. (1982) A Dictionary of Ecology, Evolution and Systematics, Cambridge University Press, Cambridge Lindroth, C.H. (1961-1969) The ground beetles (Carabidae. excl. Cicindelinae) of Canada and Alaska. Parts 16. Opuscula Entomologia, xlvii+1192pp. 1961, Part 2

Suppl. 20:l-200; 1963, Part 3, Suppl. 24:201-408, Part 4, Suppl. 29:409-648, Part 5, Supp1.33:649-944, Part 6, Suppl. 34945-1 192, Part 1, Suppl. 3ki-xlvii. Littell. R.C., Milliken, GA., Stove, W. W.. & Wolfinger, RD. (1996) SAS@System for Mixecl Models, SAS lnstitute Inc., Cery. N.C. Loreau, M. (1988) Deteminants of the seasonal pattern in the niche structure of a forest carabid community. Pedobiologiia, 31, 75-87.

Lund. R.D. & Turpin, F.T. (1977) Serologkal investigation of black cutworm larval consumption by ground beetles. Annals of the EntomoIogical Sociev of America, 70, 322-324.

Mack T.P 8 Buckman, C.B. (1990) Effeds of two planting dates and three üllage systems on the abundance of lesser cornstalk boret (Lepidoptera: Pyralidae), other selected insects, and yield in peanut fields. Journal of Economic Entomology, 88 ,1 034- t 04 1.

MacArthur, R. (1957) On the relative abundance of species. Amencan Natumlist. 94, 25- 36. MacArthur, R. (1980) On the relative abundance of bird species. Pmceedings of the National Academy of Science, Washington, 43, 293-295. Mackay, P.A. 8 Knerer, G. (1979) Seasonal occurrence and abundance in a cornmunity of wild bees from an old field habitat in southern Ontario. The Canadian Entomologist, 11 1 , 367-376.

Magunan, A. E. (1988) Ecological Divenity and its Measumment. Princeton Univenity Press, New Jersey. Mandelbrot, B.B (1977) Fractals, Fun, Chance and Dimension. W.H. Freeman, San Francisco. Manhall, CR. 8 Ward. P.D. (1996) Sudden and gradua1 Molluscan extinctions in the latest Cretaceaus of: Western European Tethys. Science, 274, 1360-1363.

May, R.M. (1975) Patterns of species abundance and divenity. Ecology and Evolufion of

Communities (eds ML. Cody & J.M. Diamond ) pp. 81-120. Harvard University Press. Cambridge. Meadows D*H+,Meadows DL., Randers J. & Behrens W.W. 111. (1972) The tirnits to Gmwth: A Report for Ihe Club of Rome's P~JWon the Pmdicament of Menkind. Signet. New York, USA. Mitchell. A. (1989) A Fragile Paradise: Netun, and man in the Pacifie. Collins. London, England. Mitchell, B. (1963) Ecology of hm> carabid beetles, Bembidion lamprof (Herbst) and Trechus quadnstn'atus (Schrank): 1. Life cycles and feeding behaviour. Journal of Animal Ecology, 32, 289-299.

Morgan, J.W. (1999) Defining grassland fire events and the response of perennial plants to annual fire in temperate grasslands of South-Eastern Australia, Plant Ecology, 144, 127-144.

Momll, W.L., Lester D.G. & Wrona A.E. (1990) Factors affecting efficacy of pitfall traps for beetles (Coleoptera: Carabidae and Tenebrionidae). Journal of Entomological Sciences, 25,284-293.

MOIT~Y,D.R. (1995) Using metal-tolerant plants to reclaim mining wastes. Mining

Engineering, 47, 247-249.

Morse, D.H. (1980) Behavioml Mechanisms in Ecology. Harvard University Press, Cambridge. Morse, R.A. 8 Flottum. K. (eds) (1997) Honey Bee Pests, Pmdaton and Diseases, 3'' ed.

A. 1. Root Company, Medina, Ohio. Motomura, 1. (1932) Patterns of species abundance and diversity . Ecology and Evolution

of Communities (eds ML. Cody & J.M. Diamond) pp. 81-120. Harvard University Press, Cambridge. Müller, J.K. (1985) Avoidance of cornpetition and niche building in carabid beetles.

ZeitschritY fur zoo/ogische Systematik und Evolutionsforschung, 23, 299-3 14. Murphy. S.D. & Swanton C. J. (1999) Responses of weeds to different faming systerns in southern Ontario: Integmted farming systems helps reduce weed pressure and weed seedbanks. Final Report January 1999. Faming Systems Research Group, University of Guelph. Neave, P. (1992) The msponse of soi1 micmflora and faune to sprnig plowing of old lFeM and zerolil sods. MSc. Thesis, University of Guelph. Nelson, W.G. (1987) An evaluation of deviation from the lognonal distribution among species as a pollution indicator in marine benthic communities. Journal of Expenmental Marine Biology and Ecology, 11 3, 18 1-208. Niemela. J.. Spence. J.R.. & Spence D.H. (1992) Habitat associations and seasonal activity of ground-beetles (Coleoptera, Carabidae) in central Alberta. The

Canadiaan Entomlogist, 124, 52 1-540.

Niemela, J. & HaIrne, E. (1992) Habitat associations of carabid beetles in fields and forests on the Aland Islands, SW Finland. Ecography, 15, 3-1 1.

Nilsson, A.N. & Soderberg, H. (1996) Abundance and species richness patterns of diving beetles (Col., Dytiscidae) from exposed and protected sites in 98 northem Swedish lakes. Hydmbiologia, 321 ,83088. Nummelin, M. (1998) Log-nomal distribution of species abundance is not a universal indicator of rainforest disturbance. Journal of Applied Ecology. 35. 454-457. Oryokoot, J.O.€., Murphy, S.D. and Swanton, C.J. (1997) Effect of tillage and corn on pigweed (Amaranthus spp.) seedling emergence and density. Weeû Science. 45,

120-126. Otto. C. 8 Svennson, B.S. (1982) Structure of communities of ground-living spiden along altitudinal gradients, Hdarctic Ecology, 5, 3547. Paoletü., MG., Schweigl, U. and Favretto, M. R. (1995) Soil macroinvertebrates, heavy metals and organochlorines in low and high input apple orchards and a coppiced woodland. Wobiologia, 39,20-33. Patrick, R. (1988) The structure of diatom communities in similar edogical conditions. Americsn Nsturalist, 102,173-183. Pausch, RD. (1979) Observations on the biology of the seed corn beetles, Stenolophus comme and Stenolophus kontei. Annals of the Entomological Society of America, 72,24-28. Pearson, O. L. & Mury, E.J. (1979) Character divergence and convergence among tiger

beetles (Coleoptera: Cicindelidae). Ecology, 60, 557-566.

Peet, N.B., Watkins, A.R., Bell, D.J., and Shama, U.R. (1999) The conservation management of Impemta cylindiica grassland in Nepal with fire and cutting: an experimental approach, Journal of Applied Ecology, 3, 374-387.

Peters. R.H. (1991) A Critique for Ecology. Cambridge University Press, Cambridge. Petenen. H. (1995) Temporal and spatial dynamics of soi1 Collembola during secondary succession in Danish heathland. Acta Zoologica Fennica, 196,190-1 94. Pielou, E.C. (1969) An Introduction to Mathematical Ecology. Wiley, New York. Pierce, B.A., 8 Sikland, N. (1985) Variation in acid tolerance of Connecticut woodfrogs; Genetic and matemal affects. Canadian Journal of Zoology, 63, 1647-1651. Pollet, M. & Desender, K. (1985) Adult and larval feeding ecology in Ptemstichus

melanadus III. (Coleoptera , Carabidae). Mededelingen van de Faculteit

Landbouwwetenschappen. 50, 58 1-594.

Pollet, M., Desender K. & van Kerckvoorde, M. (1987) Prey seledion in Lonkem pilicomis (Col., Carabidae). Acta Phytopathologica et Entomologica Hunganca. 22, 42 5- 431. Poole, R.W. (1974) An Introduction to Quanfitative Ecology. McGraw-Hill Kogakusha, Tokyo. Popova, O.A. (1978) The rok of predaceous fish in ecosystems. Ecology of fmshwater fish production. (ed. S.D. Gerking) pp. 215-249. Blackwell, Oxford.

Preston, F.W. (1962) The canonical distributions of commonness and rarity. Ecology, 43, 185-215. Preston, F.W. (1948) The commoness, and rarity, of species. Ecology, 29,254-283.

Preston, F.W. (1980) Noncenonical distributions of commonness and raflty. EcoIogy, 61, 88-97. Raukiaer, C. (1918) Recherches statistiques sur les formations vegetales. De! Danske Videnskabemes Selskab Biologiske Meddeleser 1.7-80.

Riddick, E.W. & Mills, N.J. (1994) Potential of adut carabids (Coleoptera: Carabidae) as predaton of fifth-instar codling moth (Lepidoptera: Tortricidae) in apple orchards in California. Envimnmntal Entomology, 23, 1341 -1 345.

Rivard, 1. (1964) Carabid beetles (Coleoptera: Carabidae) from agricultural lands near Belleville, Ontario. The Canadian Entomologist, 96, 51 7-520.

Roff, D. (1981) On being the right size. American Naturalist, 118, 405-422.

Roosevelt, A. (1992) Secrets of the Forest. Sciences, November-becember, 22-28. Root, R.B. (1967) The niche exploitation pattern of the bluogray gnatcatcher. Ecological Monographs, 37, 317-350. Rushton, S.P., Luff, M.L. 8 Eyre, M.D. (1991) Habitat characteristics of grasslsnd Pterostichus species (Col., Carabidae). Ecological Entomology, 16, 91- 104. Scheiner, S.M. 8 Teeri, J.A. (1987) Rapid genotypic change in a population of the gras

Denthonia spicata foltowing disturbance. Canadian Journal of Botany, 9, 1819- 1823. Scheiner, S.M. & Gurevitch. J. (eds) (1993) Design and Analysis of Ecological Expedments. Chapman and Hall, London Scheiner, S.M. (1993) Introduction: Theories, hypotheses, and statistics. Design end analysis of ecologicel expetiments. (eds SM Scheiner & J . Gurevitch) pp. 1- 13. Chapman and Hall, London Seibert, P. (1993) Vegetation and man in South America from a historical perspective. Phflocoenologie, 23,457498. Sdmann, E., Tlman, O. & Haarstad, J. (1996) Insect species diversity, abundance and body size relationships. Nature, 380, 704-708. Seimann, E., Tilmen, D. and Haarstad. J. (1999) Abundance, diversity and body size: patterns from gnssland arthropod community. Journal of Animal Ecology, 68. 824-835. Sekulid, R., Camprag, D., Kerefi, T. & Taloii, B. (1987). Fluctuations of carabid population density in winter wheat fields in the region of Backa, northeastern Y ugoslavia (196 1- 1985). Acta Phytopathologica et EntomoIogica Hungarica, 22, 265-271. Shaw, K.M., Lambshead, P.J.D. & Platt, H.M. (1983) Detection of pollution-induced disturbance in marine benthic assemblages with special reference to nematodes. Manne Ecology - Prognsss Series. 11, 795-202. Sheail, J. (1987) Semnly-fnnt yean h Ecology: The British Ecological Society. Blackwell Scientific Publications, Oxford. Steward, G.R. 8 Popp, M. (1987) The ecophysiology of mangroves: Plant life in aquatic and amphibious habitats. Speciai Publication of the Bdtish Ecological Society, 5,

333-345. Smith. C. L. 8 Taylor J.C. (1972) Results of the TeMite program: ecology of coral mef lishes. Science Bulletin, Natural History Museum of Los Angeles County.

Sokal, R.R. 8 Rohlf. F.J. (1995) Biometry. 33medn. W.H. Freernan, New York.

Southwood. T. R. E. (1978) Ecological Methods: Wifh Patticular Reference to the Study of lnsect Populations. Chapman and Hall, London. Spence. J.R. (1990) Success of European carabid species in western Canada: preadaptation for synanthropy? The deof gmund beetles in ecological and enMmnmentai studies. (ed. N.E. Stork) pp. 129-142. Anthenaeum Press, Newcastle upon Tyne. Spurr, S. H. & Bames B.V. (1980) Fomst Ecobgy. Wiley, Toronto. Stenseth, N.C. (1979) Where have dl the species gone? On the nature of extinction and the Red Queen hypothesis. OIKOS, 33,196-227. Stinner, B.R., McCartney, D.A. & Van Doren, DM., Jr. (1988) Soil and foliage arthropod communities in conventional, reduced and no-üllage corn (Maize, Zea mays L.) systems: a cornparison after 20 yean of continuous cropping. Soi1 and Tillage ReseaM, 11, 147-158.

Stubblefield, J.W., Seger, J., Wenzel, J.W. & Heisler, M.M. (1993) Temporal. spatial. sex- ratio and body-sire heterogeneity of prey species taken by the beewolf Philanthus sanbomii (Hymenoptera: Sphecidae). Philosophical Transactions of the Royal Society of London 8, 339, 397-423. Sugihara, G. (1980) Minimal comrnunity structure: an explanation of species abundance patterns. The Amencan Naturelist, 116, 770-787.

Sunderland. K.O. (1975) The diet of some predatory aithropods in cereal crops. Journal of Applid Ecology, 1 2, 507- 11 5.

Sunderland, K.D., LBvei, G.L. & Fenton J. (1995) Diets and reproductive phenologies of the introduced ground beetles, Hatpalus amus, and Clivina australasMe

(Coleoptera: Carabidae) in New Zealand. Austmlian Journal of Zoology, 43, 39-

50. Sustek, 2. (1981) Influence of dear cutting on ground beetles (Coleoptera. Carabidae) in a pine forest. Communicationes Instituti Forestalis dchoslovaniae .1 2, 243-254. Swanton, C. J. & Weise, S.F. (1991) lntegrated weed management: the rationale and approach. Weeâ TechnoIogy, 5,657-663. Taylor, L.R., Kempton, R.A. 8 Woiwod, I.P. (1976) Divenity statistics and the log-series modal. Journal of Anime1 Ecology, 45,255272.

Thomas, M.R. & Shattock, R.C. (1986) Filamentous fungal associations in the phylloplane of Lolium penenne. Tmnscnpts of the British MycoIogical Society, 87, 255-268. Tilman, O. & Downing. J.A. (1994) Biodivenity and stability in grasslands. Nature, 367, 363-365. Tepedino, V. J. 8 Stanton, N.L. (1981) Diversity and competition in bee-plant communities on short-grass prairie. OIKOS, 36, 35-44.

Thiele, H-U. (1977) Carabid Beetles in their Environments: A Study on Habitat Selection

by Adaptations in Physiology and Behaviour. Zoophysiology and Ecology. vol. IO. Springer-Verlag, Berlin. Tokeshi, R. (1993) Species abundance patterns and community structure. Advances in Ecological Research, 24,111- 195. Tolonen, T. (1995) Importance of generalist epigeal predator species in a cereal field:

predation on baits. Journal of Applied Entomology, 11 9.1 1 3- 11 7.

Tonhasca, A. Jr. (1993) Carabid beetle assemblages under divenified agroecosystems. Entomologia Experimentalis et Applicata, 68,279-285.

Tonhasca, A. Jr. & Stinner, B.R. (1991) Effects of strip intercropping and no-tillage on some pests and beneficial invertebrates of corn in Ohio. Envimnmental Entomlogy, 20,1251-1258.

Turner, R.M. & Brown, D.E. (1982) Desert Plants, University of Arizona, Tucson.

Tsuyuzaki, S. (1994) Fate of plants from buried seeds on volcano Usu, Japan, after the 1977-1978 eruptions. Amencan Journal of Botany, 81,395-399. Tyler, B.M.J. & Ellis, C.R. (1979) Ground beetles in three tillage plots in Ontario and observations on their importance as predators of the northern corn rootwom, Diabrotica longicomis (Coleoptera. Chrysomelidae). Proceedings of the Entomological Society of Ontario, 11 O, 6573.

Ugland, K.I. & Gray, J.S. (1982) Lognomal distributions and the concept of community equilibrium. OIKOS, 33, 196-227. Vandenneer, J., Brenner, A. 8 de la Cerda. I.G. (1998) Growth rates of tree height six yean after humcane damage at four localities in eastern Nicaragua. Biotopiica, 30, 502-509. Varis, A-L. (1989) Cabbage field Carabidae (Coleoptera) and their role as natural enemies of Delia radicum and O. florelis (Diptera, Anthomyiidae). Acta EntomoIogica Fennica, 53, 6 1-63. Vickerman, G.P. & Sundedand, K.D. (1975) Arthropods in cereal crops: noctumal activity, vertical distribution and aphid predation. Joumal of Applied Ecology, 12, 755-766. Ward, P.S. (1973) The eîfects of environmental disturbance on species diversity and species-abundance patterns in a taxocene of phytophagous insects. Report for Biology 537, Queens University, Kingston Ontario. Whittaker, R.H. (1965) Dominance and divenity in land plant communities. Science, 147, 250-260. Whittaker, R. H. (1972) Communities and €cosystems+Macmillan, New York. Whittaker, R.H. (1977) Evolution of species diversity in land wmmunities. In Evolutionary Biology, Vol. IO(eds M.K. Hecht, W.C. Steere & B. Wallace) pp. 1-67, Plenum, New York Wilson, E.O. (1998) Consilience: The Unity of Knowledge. Knopf, New York. Wilson, J.B. ((1993) Would we recognize a broken-stick community if we found one? OIKOS, 67,181-183. Wisser, S.K., Allen, RB. & Platt, K.H. (1997) Mountain beech forest succession after a fire and Mount Thomas Forest, Canterbury, New Zealand. New Zealand Journal of Botany, 35, 505-5 15. Witman, J.D. (1992) Physical disturbance and cornmunity structure of exposed and protected reefs: A case study from St. John, U.S. Virgin Islands. American Zoologist, 32,841654. fou, X., Zucca, C. P. Waide, R .B. 8 MCDOW~II~W. H. (1995) Long-terrn influence of deforestation on tree species composition and litter dynarnics of a tropical rain forest in Puerto Rico. Forest Ecology and Managemsnt, 78, 147-157. APPENDIX 1: ECOLOGY OF CARABID BEETLE SPECIES COLLECTED FROM SOU1HERN ONTARIO FARM FIELDS

1. AGONUM Bonelli

Agomm cupreum Dejean This 7.0-9.5 mm long Agonum species is transamencan and can be found in open country with sparse vegetation (Lindroth 1981-1969). In the field the species was observed to eat Hylemya brassicae (Bche) and ladybugs, and in captivity it was observed to eat eggs of Euxoa ochrogaster (Guén.) (Lepidoptera: Noctuidae) (in Larochelle 1990).

Agonum cuprlp.nne Say This 7.5-9.3 mm long Agonum species can be found across North America where there is moist soi1 and habitats ranging from open ground to dense vegetation (Lindroth 19614969). In the field the species was observed to eat crickets and Hylemya antique

Meig. (in Larochelle 1990). In captivity it was observed to eat maize, ham. earthwoms, chicken, and a dead Agonum placidum Say (in Larochelle 1990).

Agonum &rl#n#tonl Lindroth A. darlingtoni, a 4.9-5.9 mm long hygrophilous (i.e. are attracted to water) species (Lindroth 1961 -1989), is found in eastem Canada and north-eastem U.S.A. (Lindroth 1961 -1969). The feeding habits of the carabid have not been well established. Agonum gratiosum Mannerheim A. gmtiosum tend to be found on open, usually moderately moist ground (Lindroth 1961 -1969). The beetles are 6.4-9.1 mm long. This Agonum is native to North America and is distributed across the continent (Lindroth 1961 -1969). In captivity the species ate meal woms (Tenebrio spp.), and pieces of ham (in Larochellel990).

Agonum hypolithos Say This 13.5-15.5 mm long species has a distribution which is bordered by Pennsylvania to the south, Michigan to the west and southem Ontario to the north (Lindroth 1961 -1969). It tends to be found beneath Stones in moist habitats (Lindroth 1961 -1969). The feeding habits of A. hypolithos are not well known.

Agonum mue/lerl Herbst A. muellen' is a Palaearctic species, which was introduced to both coasts of North America (Lindmth 1961 -1969). This 7.2-9.5 mm long (Lindroth l96l-l989) beetle has been recorded in open, moderately dry fields (Lindroth 1961-1969; Thiele 1977). In the field the beetle was observed to feed upon wireworm (Agdotes sputator) (Erwin et al. 1976) In captivity the species was observed feed upon beef, small larvae of insects and small woms (in Larochelb1990).

Agonum placidum Say A. placidum is a xerophilous species that has been recorded on cultivated soi1 arnong weeds (Lindroth 1961 -1969; Rivard 1984). This 6.8-8.8 mm long species has a transarnerican distribution, but it does not reach Me Pacific coast (Lindroth 1961 -1989). In captivity the species was observed to feed upon Euxoa ochmgaster Guen. eggs and adults (Red-backed cutworm) (Erwin et al.1976; Frank Ig71a), Wheat midge larvae (Floate et al. 1990). Additional food items for the species in captivity recorded in Larochelle (1990) are a maire kemel, pieces of ham and a slug. In the field the species was observed to eat C. pomonelle larvae (Riddick 8 Mills 1994).

Agonum plccolum Leconte

This transamerican species is 6.0-7.6 mm long (Lindroth 1961 -1969). It occurs on bare or spanely vegetated soi1 (Lindroth 1961 -1969). The feeding behaviour of the species is not well established.

Agonum sordens Kirby A. sordens is a transamerican species (Lindroth 1961 -1969). 00th Lindroth

(1961-1969) and Rivard (1964) state this carabid species prefers moist habitats, however according to Lindroth it favoun areas with moderately dense vegetation whereas Rivard suggests that the species is found on open ground with sparse vegetation. Feeding habits for the 5.3-6.5 mm long carabid have not been documented.

Agonum sulcipenne Hom There is very little documentation on the ecology of this 7.0-8.0 mm long carabid (Lindroth 1961 -1969).

Agonum mue Leconte Agonum tenue is a 8.0-9.5 mm long carabid whose native distribution is eastern North America. It has been recorded in manhes and at the margin of ponds, on soft, wet soi1 Wth organic content and rich vegetation, e-g.. Typha letiîolie and Carices (Lindroth 1961 -1969). Feeding habits of this beetle have not been documented. Aman aenea de Geer A. aenea is a European introduction that is restncted to northeastem North

America (Lindroth 1961 -1969). It is found on dry, open grassland (Lindroth 1961 -1969). Hagley et al. (1982) indicate that this 6.2-8.8 mm long species is polyphagous. In captivity the beetle was observed to feed upon apple maggot pupae, chickweed and dandelion seeds (Hagley et al. 1982). Davies (1959) recorded that the species ate plant material.

Aman avlda Say This 7.4-9.8 mm long species is almost transamerican, but does not mach the Pacific Coast (Lindroth 1961 -1969). According to both Rivard (1964) and Lindroth (1961 - 1969) A. avida is found in open ground with moderate to dense vegetation. In both the field and Iaboratory, it was observed to feed upon E. ochmgaster (Frank 1971a; Erwin et al. 1976).

Amam carhata Leconte This 11.O-14.2 mm long species occun in the central part of North America, and can be found on open, moderately moist grassland areas (Lindroth 1961 -1969). In the field. the species has been observed to feed upon Grarnineae, seeds and fungi (P8mnospom sp.) (in Larochelle 1990). Amaru convexa Leconte A. convexa is a xerophilous species that is confined to dry. open grassland (Lindroth 1961 -1969). According to Lindroth (1961-1969) the distribution of this carabid is almost transamerican, and occurs south to New Mexico. lndividuals are 5.3-8.0 mm long (Lindroth 1961 -1969). In the field the species was observed to feed upon ladybugs (Lamchelle 1990).

Amara cupmoIaîa Putzeys This 6.5-9.0 mm long species has a transamerican distribution. It has been found in sandy gravel pits with spane vegetation (Lindroth 1961 -1969). In the field the species has been observed to feed upon grass seeds ( Johnson & Cameron 1969; Erwin el al.

1976), alfalfa weevil larvae (Bamey & Pass 1986b), weed seeds (Barney & Pass 1988b), and amyuvomis (Clark et al. 994) .

Amara familieris Duftschmid This 5.6-7.2 mm long species was introduced into North America (Lindroth 1961 - 1969). A. familiaris is found near hurnan habitations in open ground with moderate to dense vegetation (Lindroth 1961 -1969; Rivard 1964). In the laboratoiy this carabid has been observed to feed upon grass seeds (Johnson & Cameron 1969, Emin et al. 1976), second-instar amyworrns (Clark el al. 1994), and Cemstium species, seed, flowers (Davies 1959), barley and winter wheat (Vickeman & Sunderland 1975). In the field the species was observed to feed upon aphids (Vickerrnan & Sunderland 1975). Amwa ktior Kirby Both Erwin el al. (1976) and Frank (1971 a) report that A. latior consurned Eochmgaster under labonitory conditions. Larochelle (1990) notes in the field that egg- pods of locusts (Orthoptera) were eaten. The native distribution of this 7.8-10.8 mm long carabid is transamerican (Lindroth 1961 -1989). Rivard (1964) found this species on open dry ground, and Lindroth (1961-1 969) recorded this Aman species on sandy soil.

Amara littoralis Mannerheim A. littomlis has a transamerican distribution (Lindroth 1961 -1969). The 6.2-9.3 mm (Lindroth 1961 -1969) beetle can be found in open, moderately dry areas with rich weedy vegetation (Lindroth 1961 -1969). In captivity the beetle was observed to feed upon a dead fly, pieces of ham and apple. In the field the beetle was observed to feed upon egg pods of Camnula and Melanoplus (Otthoptera; Acrididae) (in Larochelle 1990).

Amm rubricr, Haldeman There is little documentation of the ecology of this 5.2-7.3 mm long carabid (Lindroth 1961 -1969). However Lindroth (1961-1 969) does mention Mat he detected it abundantly in a gtavel pit on a sandy moraine. As well, the species was seen to feed upon flowers of Solidago (Compositae) (in Larochelle 9990).

3. ANISOOACTYLUS Dejean

Relatively little has been documented about this 9.2-10.8 mm long beetle (Lindroth 1961 -1989). Anlsod~ctyluswrticur Say AIthough common in eastem Norîh Amena, the western distribution of the 8.8- 11.O mm long beetle is not known (Lindroth 1961 -1969). The habitat preference of the species is dry, open ground with thin to dense vegetation ( Rivardl964; Lindroth 1981 - 1969). ln captivity, A. msficus was observed to feed upon grass seed (Johnson & Cameron l969), maire kemels (in Larochelle 1990), pieces of earthwonn (in Larochelle 1990) and a slug (in Larochelle 1990). In the field, animal food made about one-fourth of total diet for this species and the vegetative portion of their diet was denved from June grass and other graminaceous plants (Johnson & Cameron 1969).

AnIsOdactylus sunctaecrucls Fabricius This 8.340.5 mm long beetle has a transamerican distribution however, its southern limit is not known (Lindroth 1961 -1969). The species is almost hygrophilous and has been found to occur near water amongst dense vegetation (Lindroth 1961 - 1969). The species is polyphagous. In the field, it was observed to feed upon insect fragments, animal food, vegetation and grass seeds (Erwin el al. 1976). Under laboratory conditions, Baines et al. (1990) report the species consumed carrot weevils and carrot seeds. Hagley et al. (1982) found that the insects feed equally well on weed seeds, apple maggot pupae and codling rnoth eggs, larvae and pupae. It also eats early instars of C. pomonella (Riddick 8 Mills 1994).

4. BEMBlOlON Latreille

BembMion rnmicunum Dejean There is relatively little recurded about this 5.06.0 mm long beetle. However Lindroth (1961-196389) notes it can be found in eastem North America. BembMion Inaequale Say This 4.6-5.6 mm long species is found in eastern North America (Lindroth 1961 - 1969). The species prefers moist soil with sparse vegetation (Rivard 1964; Lindroth 1961-1969). Feeding habits of the beetle have not been recorded.

Bembidion levettei Casey

This 5.4-6.3 mm long species has a transamerican distribution. The species prefers moist soi1 with spane vegetation ( Lindroth 1961-1969). Feeding habits of the beetle have not been recorded.

Bembidion nkidum Kirby Although Lindroth (1961-1969) States that B. nitidum is a xerophilus species, Rivard (1964) found the species prefen moist soils. The 4.1-5.3 mm long species has an almost transamerican distribution (Lindroth 1961-1969). In the field it was observed to eat cabbage root fly eggs (Thiele 1977). In captivity it was observed to feed upon E. ochmgaster eggs (Erwin et al. 1976; Frank 1971a). Additional food items eaten by the species in captivity include eggs of Hylemya bmssicae @ch&),a maize kemel (in Larochelle 1990), pieces of ham (in Larochelle 1990) and earthwoms (in Larochelle 1990).

8embidIon oûetîhuri Hayward B. obsrthud occun rnainly in eastem North America (Lindroth 1961-1969). The 4.04.5 mm long species occun near wming water in areas with bare vegetation (Lindroth 1961 -1969). The feeding habits of this carabid have not been well documented. Bembidion obscurellum Motschulsky Both EnMn et al. (1976) and Frank (1971a) report that under laboratory conditions this carabid consumed E. ochrogaster eggs. In the field ladybugs were eaten (in Larochellel990). The species is 3.9-5.1 mm long and has a circumpolar distribution (Lindroth 1961 -1969). It is often found near water in areas with spane vegetation (Lindroth 1961 -1969).

Bembidion obtusum Serville This 2.8-3.5 mm species is a recent European introduction (Lindroth 1961 -1969). It is found mainly in cultivated fields (Lindroth 1961 -1969; Thiele1977). Food items consumed in the field include Collembola, Acarina. spores, pollen and vegetal tissue (in Larocheile 1990).

Bembidion quadrimaculiturn oppasitum Say B. quadhaculatum oppositum is a 2.8-3.7 mm long carabid with a Paleoardc distribution (Lindroth1961-1969). It has been found in a wide vanety of habitais ranging from dry open ground (Rivard 1964; Floate et al. 7990) to lake shores and river banks (Lindroth 1961 -1969). This species is common in cultivated areas (Lindroth 1961 -1969; Thiele 1977). The Bembidion is omnivorous. In the field the species was observed to feed upon strawôerry leaves (Erwin et al. 1976), and cabbage root fly (Delia radicum) eggs (Thiele 1977). In captivity the beetle was observed to feed upon fruit fly pupae (Tolonen 1995), wheat midge larvae (Floate et al. 1990), carrot weevil (preferably pupae) and canot seeds (Baines et eLl99û), black cutwom eggs (Best & Beegle 1977a), root maggot (Delia spp.) eggs and larvae (Grafius & Wamer 1989), com rootworm eggs (Grafius & Wamer 1989), E. ochrog8stereggs (Frank 1971a) and 0. radicum (Vans 1989). Bembidion rripldum Leconte This 3.84.4 mm long species is often found near bodies of water (Lindroth 1961 - 1969). Its native distribution is east of the Rockies, but it does not reach the Atlantic coast (Lindroth 1961 -1969). Under laboratory conditions it was observed to feed upon black cutwomi eggs and larvae (Best & Beegle 1977b, Baines et al. 1990).

Bembidion ruplcol. Kirb y Bath Erwin et al. (1976) and Frank (1971a) report that under laboratory conditions this carabid was obsetved to feed upon E. ochmgestereggs. According to Lindroth (1961-1969) the species is pronouncedly xerophilous and occun on cultivated fields. The 4.1-5.5 mm long species has a distribution that in the late 1960's was found in central North America (Lindroth 7961-1 969).

Bembidion sslebratum Leconte B. salebratum is found east of the Rockies (Lindroth 1961 -196s). The species is usually found near water in areas with tittle vegetation (Lindmth 1961 -1969). Food preferences of the 3.7-4.9 mm long beetle haven' t been documented (Lindroth 1961 - 7969).

BembidIon sordldum Kirby This species is transamerican in its distribution. The 5.2-7.3 mm long species is found on moist clayish soi1 with spanr vegetation. Food preferences of the beetle has not been documented (Lindroth 1961 -1969) 1 Bembidion tetraco/um Say 6. tetmcolum is a west-Palaearctic species that was introduced into North Arnerica (Lindroth 1961 -1969). The 4.9-6.1 mm long beetle is found on open ground with moist soi1 and sparse vegetation (Rivard 1904) and often on cultivated fields (Lindroth 1961 -1969). Thiele (1977) notes that the species eats mainly srnall mites, pseudoscorpions, small insects, Coltembola and insect eggs.

Bembidion tnnsvenate Dejean This 6.1-7.9 mm long species has a transamerican distribution (Lindroth 1961 - 1969). t is found on barren grave! (Lindroth 1961 -1969). Its feeding behaviour is not well recorded. Bembidion vemico/or Leconte This Bembidon has a transamerican distribution, although it is predominately northem (Lindroth 1961 -1969). The 2.8-3.6 mm long species is found on open ground with moist soils and sparse vegetation (Lindroth 1981 -1969. Rivard 1964). In captivity B. mr~icolorwasobserved to feed upon E. ochrogaster eggs (Frank 1971; Emin et al. 1976).

5. CALATHUS Bonelli

This large (7.0-1 1.1 mm long) carabid has a transamerican distribution (Lindroth 1961 -1969). Lindroth (1961 -1 969) reports that it can be found among dead leaves and rnoss under bushes and deciduous trees on either moist or dry ground. In captivity the species was observed to feed upon freshwater shfimps (in Larochelle 1990). 6. CALLEIDA Dejean

CaIIeida punetata Leconte The adults of C. punctata are specialized predators of Noctuid and Pyralid moth caterpillars (Lindroth 1961 -1969). Lindroth (1961-1 969) did not note the habitat preferences of this 7.0-8.5 mm long carabid. Its distribution is in east south to St. Louis, west to Kansas and Michigan, north to Ottawa (Lindroth 1961-1969).

7. CALOSOMA Weber Calosoma calidum Fabricius According to Erwin et al. (1976) in captivity this 19.0-25.0 mm long species was obsewed to feed upon Lepidoptera larvae Katydids, Stone crickets, stinkbugs, cicada nymphs, ants, sowbugs, jumping spiders, buttetfly chrysalis, rneasuring Worms, Noclua clandestina, Potthetna dispar (Gypsy moth), Malacosorna amen'canum (Eastern Tent caterpillar), Hyphantnacunea spp. and E. ochmgaster. See Latochelle (1990) for additional food items. It is widely distributed in North America and is found in open, rather dry fields with low vegetaüon (Lindroth 1961 -1 969).

8. CHLAENIUS Bonelli

Chlaenius lithoph~lusSay The distribution of C. lithophilus is almost transamerican, but it does not mach the Pacific coast (Lindroth 1961 -1989). In the laboratory this 8.0-9.5 mm long beetle was obsewed to feed upon earthwons, dead slugs, maize kernels (Erwin et al. 1978). The species is found aî the border of ~nningand standing. often small bodies of water, where vegetation is rich, but also on soft mud (Lindroth 1981 -1969), and is often associated with C. sericeus or C. tricdor (Lindroth 1961 -1989).

Chlaenlu. tricolor Dejean This species has been documented to occur near running water, flood plain forests and open ground with moist soi1 (Lindroth 1961 -1969; Rivard 1964). The 1O. 1- 13.5 mm long carabid has an almost transamerican distribution (Lindrothl969). Under labontory conditions it was observed to eat dead Chlaenius sericeus, dead slugs, and earthwoms (Erwin et al. 1976).

9. ClClNDELA Linné

Ciclndela finnosa grnerosa Dejean This large (10.0-18.0 mm long) species inhabits dry, spanely vegetated, sandy amas and is found on bare sandy spots and along sandy paths and roads (Dillon & Dillon 1961; Graves 1965; Boyd 1978). The species is predaceous (see Larochelle 1990).

Cichdela punctulata Olivier

C. punctulata is found on wide range of habitats but prefen dry, hard-packed sand or sandy loam (Boyd 1978). This 11 .O-14.0 mm long (Dillon 8 Dillon 1961) species had been recorded to be an opportunistic frugivore (Hill & Knisley 1992), however according to Larochelle (1990) the species is predaceous.

Cicinakla np~ndaOejean

Although this Cicindela crin k an opportunistic fnigivore (Hill & Knisley 1992) it is mainly pndaceous (in Larochelle 1990). The 12.0-13.0 mm long (Dillon & Dllon 1961)

134 beetle can be found in a wide variety of habitats, from open sandy areas, to borden of fields and dirt roads, shores of lakes to mud flats (Boyd 1978).

Cicindela scutetluris lecontei Haldeman This 12.0-13.0 mm long species (Dillon & Dillon 1961) is found dry, sandy areas (Graves 1965). The species is predaceous (in Larochelle 1990).

Cicindela sexeutfata Fabricius This 10.0-16.0 mm long species can be found in forests, along paths, roads, bkw oub and other open areas (Kaulban 8 Freitag 1993), as well as pastures near wooded areas (Boyd 1978). The distribution of the species to the south and east is limited by Gulf of Mexico and Atlantic Ocean (Boyd 1978). The species is predaceous (in Larochelle i990).

10. CLlVlNA Latreille

Ctivinrr mer Linné C. fossorwas introduced from Europe in the 1960's (Lindroth 1961 -1969). It is found on cultivated soils parks and gardens (Lindroth 1961 -1969). The 5.5-6.5 mm long beetle is omnivorous (Forsythe 1983). In the field it has been observed to eat cereal aphids (Rhopalosiphum padr)(Sunderland et a/. 1995), Agdotes sputator (Erwin et al. l976), strawbemes (Lindroth 196 1 -1969) and other crop plants (Desender et al. 1985; Vans 1989). ln captivity, C. fossor was obsewed to feed upon larvae and pupae of blossom beeUe (Meligethes sp.) (Thiele 1977), canot weevil (Listronotus oregonemis) eggs larvae and pupae (Sunderland et al. 1995), canot seeds (Sunderland et a/.1995) wireworms. (Sunderland et al. 1995) and tumip rwt fly eggs (Sunderland et al. 1995) 11. COLLlURlS De Geer

CoIlIurus pensylvanica Linne Lindroth (1961-1969) recorded that this 5.8-7.2 mm long carabid occurs south to Mexico. west to Califomia, nom to Grand Bend, east to Montreal. Its preferred habitat is open grassland with sparse vegetation (Lindroth 1901 -1969; Rivard 1964). In the field, the species was observed to feed upon lanrae of Ancylis comptana fiagada W. and R. (Lepidoptera: Tomicidae) and Blissus leucopterus Say (Heteroptera: Lygaeidae) (in Larocelle IWO).

Diplocheila obtusa This 9.7-1 1.7 mm long carabid has a transamencan distribution with a gap in central Canada (Lindroth 1961 -1969). Its habitat preferences are not well recorded. In captivity it was obsenred to feed upon pieces of earthwonns, ham, maire kemels and a raisin (Larocheils 1990).

13. DYSCHlRlUS Bonelli

Oyrchlrius bnvispinu~Lecon te There is little published information on the 3.4-4.2 mm long beetle (Lindroth 1961 -1969). Oyrehirlur ~rylhrocefusLeconte The 4.44.9 mm long beetle is found near bodies of water. b distribution extends frorn Florida to Ohio to southem Ontario and to western Newfoundland (Lindroth 19û1 - 1969). The feeding habits of D. erythmnrs have not been well recorded.

Dyachirius globuIosus Say Thiele (1977) noted that this small(2.6-3.2 mm long) insect hunts staphylinid beetles. The beetle has an almost transarnerican distribution, but not reach the Pacific coast (lindroth 1961 -1969). D. globulosus has been found in cultivated fields (Thiele 1977), as well as open ground with sparse vegetation (Rivard 1964) and near river banks

(Lindroth 1961 -1969).

Dyschirius p/îtus Dejean This 3.4-4.9 mm long carabid has a distribution that is both circumpolar and transamerican (Lindroth 1961 -1969). The species can be found in open areas among low scattered vegetation (Lindroth 1961 -1969). In the field the species was observed to feed upon Bledius fuscipes (Coleoptera: Staphylinidae) (in Larochelle 1990).

4. GEOPINUS Leconte

Geopinus Incrassatus Dejean In the field this large (13.0-17.0 mm long) beetle was observed to feed upon confer seedlings (Johnson & Camemn 1969), cutwons (in Larochelle 1990). and seedlings in seed beds (in Larochelle 1990). As well. in captivity it was observed to feed upon Musca domesfica L. (Diptera: Muscidae) (in Larochelle 1990). It is found in the eastem half of North America (Lindroth l98l-l989). Rivard (1964) stated G. incmssatus is found on open, dry ground with moderate to dense vegetation, however according to Lindroth (1981-1969) the species prefen areas with sparse vegetation.

HARPALUS Latreille

HarprrIus a#nla Schrank H. alKnis can be found in open habitats that have moderate to dense vegetation (Lindroth 1961 -1969; Rivard 1964). The species is an omnivore, although primaflly has a vegetarian diet (Sunderland et al. 1995). In the field Sunderland et al. (1995) observed that it was observed to feed upon seeds of Stellana, Bmssica spp., chickweed. dandelion, crab grass, knotweed, red clover and strawbeny as well as Aphididae. Diptera, Nematoda, Lumbricidae. Enchytraeidae, Araneae, Coleoptera, Lepidoptera., A. sputator, immature stages of cabbage root fly (Deliaradicurn), pupae of apple maggot fly

(Rhagoletis pomonella) and aphids. Holliday & Hagley (1978) also reported that H. aliinis was predominantly phytophagous but consumed Lepidoptera and Diptera laivae. Erwin et al. (1976) mention only that the species is a voracious grass and seed feeder. Under laboratory conditions H. amis was observed to feed upon eaily instan of C. pomonella (Riddick & Mils 1994, earthwoms (Sunderland et al. 1995), and eggs, larvae and pupae of C. pomonella (Sunderland et al. 1995), pupae of apple maggot fly (Sunderland et al. 1995), eggs of cabbage root fly (Sunderland et al. 1995) and of tumip root fiy (Sunderland et al. 1995), and al1 instars of the cereal aphid Sitobion awnae (Sunderland et el. 1995). Hagley et el. (1982) noted that this carabid feeds equally well on weed seeds, apple maggot pupae and codling moth eggs, larvae and pupae. Harp.1~8bicolor Fab ricius The 10.7-14.2 mm long beetle is found in open country, often on sandy fields where the soi1 isn't too dry (Lindroth 1961-1 969) throughout eastem North America (Lindroth 1961 -1969). Under field conditions the beetle was observed to eat Blissus leucoptefus Say (Heteroptera: Lygaeidae) and Nysius angustatus Uhl. (Heteroptera:

Lygaeidae) (in Larochellel990).

Harpalus caliginosus Fabricius The 17.5-25.5 mm long beetle has a transarnerican distribution (Lindroth 1961 - 1969). In the field, the carabid fed on Amy woms, grasshoppers, stnwbenies, Ambmsia spp. seeds, caterpillan, Diptera, mites, animal food, cankerwoms, grass tissue, pollen and fungi (Erwin et a1.1976). Under laboratory conditions this species reported was observed to feed upon grass seed (Johnson & Cameron 1969; Erwin et al. 1976), gypsy moth larvae (Erwin et al. 1976), inch woms (Erwin et al. 1976), tree crickets (Oecanthus)(Erwin et a/.1976), ant pupae (Ewin et al. 1976) and earthworms (Erwin et al. 1976). The species is a serious strawberry pest in Europe and US (Erwin et al. 1976). Hatpalus emticus Say H. enaticus is found east of the Rockies, south to Alabama and New Mexico, east to Quebec and up to the North West Territories (Lindroth 1961 -1 969). This 11.O-18.0 mm long can be found in dry and open areas (Lindroth 1961 -1989). In the field the species was observed to feed upon grass seeds and Cdomfis asparagi L. (Coleoptera: Chrysomelidae) (in Lamchelle 1990). tîarplus erythmpus Dejean This carabid is found on fields and in open woods (Lindroth 1961 -1969). The distribution of the 9.7-11.4 mm long beetle is south to Florida, west to Colorado, east to Nova Scotia and north to Manitoba. Larochelle (1990) notes that in the field, the species was observed to feed upon larvae of Phyllophaga sp. (Carabidae: Scarabaedae) and in captivity the species was observed to feed upon prepupae and pupae of Altica chalybeea III. (Coleoptera; Chysornelidae).

Harpalus faunus Say The distribution of this 8.4-13.0 mm long beetle is south to Louisiana, west to Arizona, east to Terrebonne Quebec and north to Lake Huron. Lindroth (1961-1969) recorded it in a grave1 mine. In the field it was observed to eat larvae of Cmmbus caliginosellus Clem. (Lepidoptera: Pyralidae) (in Larochelle 1990).

Harplus funerwius Csiki This 10.3-13.0 mm long species found east of the Rockies (Lindroth 1961 -1969). It can be found on open, dry soi1 with sparse vegetation. In captivity it was observed to feed upon eggs and larvae of E. ochrogsser (in Larochelle 1990).

HatpIus herbivaeus Say This 9.0-1 1.O mm long beetle has a transamerican distribution, but is scarce in the prairie provinces (Lindroth 1981 -i969). The species can be found in open habitats (Lindroth 1961 -1969). H. henbi~guseats animal food, insects, Dipten. vegetable matter, fungi, grass seed, young shoots of Poe pmtenJis (Erwin et a/.1976). Also see

Laiochelle (1990) for a list of the food eaten by this species. Hatplus longtcollts Leconte The 10.3-13.5 mm long species is found south to the Gulf coast, west to South Dakota and Kansas and up to Ontario and Quebec (Lindroth 1961 -1969). Lindroth (1961-1969) found H. longicollis in a grave1 pit and in damp grassy habitats. In the field, the species was obsewed to feed upon Phyllophaga anxia LeC. (Coleoptera: Scarabaedae) and Popilliejeponica Newrn. (Coleoptera: Scarabaedae) (in Larochelle

1990).

HatpuIus pensy~vanlcusDe Geer H. pensylvanicus is a 10.1-15.2 mm long beetle that is widely distributed across North America (Lindroth 1961 -1969; Kirk 1973). It is often found in cultivated fields (Lindroth 1961 -1969; Kirk 1973), grassy areas (8nist et al. 1986) and dry, open areas (Rivard 1964). Feeding behaviour of the beetle has been described as facultatively phytophagous (House & Al1 1981). Under labofatory conditions the beetles was obsewed to feed upon inch worms (Erwin et al. 1976)' Asparagus beetle eggs (Cnocetis asparag~] (Emin et al. 1976), corn root wom larvae (Diabmtica viricifom) (Erwin et al. 1976), adult weevils (Bamey & Paso 1986), lepidopteran larvae (Bamey & Pass 1986). crabgrass seeds (8amey & Pass 1988). 5th-instar C. pomonella (Riddick & Mills (1994), 4th-stage blacû cutwom larvae (Best & Beegle 1977a. b), seeds (Best & Beegle 1977a), crickets (Best & Beegle 1977a). green cloverwom larvae (Best 8 Beegle 1977a). H. pensylvanicus eggs (Kirù 1973), and Ist-stage larvae and eggs of western corn rootmxms (KiM973). In the field, the beetles were observed to eat the following: ragweed seeds and pollen (Ambmsia ariemisiaefolia)(Lindroth 1961 -1969; Erwin et al. 1976), wheat kemels ( Kirk 1973; Erwin et al. 1W6), timothy seeds (Kirk 1973; Erwin et al. 1976). panic grass seeds (Panicum crugalli)(Erwin et al. 1978), animal food (Erwin et al. 1976). ant mites (Erwin el al. 1976), cankeimmns (Erwin et al. 1976), June grass (Erwin et al. 1W6), fun@ (Erwin et al. 1970), honey bees (Kirk 1973; Erwin et al. 1976),

17. MICROLESTES Schmidt-Goebel

MlcmIestes brevilobus The ecalogy of the 3.0-3.4 mm long carabid has not been well documented.

Notobl. plceus (lormerîy Anisotmus plceus) The food eaten by this 9.0-1 1.O mm long beetle has not been docurnented. It is common on dry, sandy fields (Lindroth 1961 -1969), and is found in the eastern part of North America (Lindroth 1961 -1969).

Notiobia termnsta (fomerly Anisofanus terminatus) The habitat preference of this 8.5-9.1 mm long beetle is dry, open, sandy ground with spane vegetation, for instance on cultivated land and in grave1 pits (Lindroth 1961 - 1969). N. teminata is found in eastern North America (Lindroth 1961 -1969). It is a very active grass feeder both in the field and laboratory (Emi'n et a1.1976). Lindroth (1961- 1969) observed that the beetle fed on seeds of fireweed (Emhtltes hiereciolia). In captivity the species was obsenred to feed upon Coleoptera without legs, caterpillan and pupae of Lepidoptera (in Larochelle 1990).

Notiophi/us remeus Herbst AIHiough Rivard (1964) mported that this species is found on dry soi1 in or near woods, according to Lindroth (1961-1969)the beetle occum in moss or amongst dead leaves in damp (but not too wet) places in deciduous woods. The 5.0-5.7 mm long beetle is found in eastem North America (Lindroth 7961 -1989).

20. OMOPHRON Latreille

Omophrw, amrricrnus Dejean This 5.1-7.0 mm long carabid is widespread throughout North America (Lindroth 1961 -1969). It is found on bare sand or sand-mixed clay, usually at standing waters, sometimes quite small pools in sandpits (Lindroth 1961 -1969). In captivity the beetle was observed to feed upon meahvomis and pieces of ham (in Larochelle 1990).

OxypseIaphus pusIIIu8 (fomeily Agonum punctlceps Casey) 0.pusillus is a transamerican carabid that can be found arnong dead leaves, in shady places and usually near water (riven and lakes) (Lindroth 1961 -1989).

21. PATROBUS Dejean

PIlrobus longlcomls Say This transamerican carabid is 9.2-14.8 mm long (Lindroth 1961 -1969). It can be found in a variety of habitats including areas near lakes and nven, rneadows, light deciduous forests and cultivated ground (Lindroth 1961 -1969). Rivard (1964) found the species on moist. open ground with moderate to dense vegetation. In the field this wrabid was observed to feed upon animal food, vegetable matter, caterpillars, aphids and grass (Etwin et al. 1976). 22. POECILUS (fomerly part of PTEROSTICHUS)

Poecilus chulchSay (fomerly known as PlerOaIchur ch8lcifw Say) This 10.5-13.0 mm long carabid can be found both on fields and in open woods, on moist soi1 (Lindroth 1961 -1969; Kirk 1975b). It occurs in south to Louisiana and west to South Dakota and Nebraska, in southern Quebec and southern Ontario residents (Lindroth 1961 -1969, Kirk 1975). Emin et al. (1976) record that in the field the beetle was observed to feed upon insect fragments, and Lund & Turpin (1977) state that it consumed black cutworm lawae. In the laboratory P. chalcites was observed to feed upon June beetles (Erwin et al. 1976). Oiabrotica bdtaeata (Erwin et al. 1976), western corn rootwotm (Diabmtica viqifera) eggs, larvae and adults (Kirk 1975), second- and fourth-instar arrnywoms (Clark et al. 1994, Black cutworm eggs and 4th-stage larvae (Best 8 Beegle1977b), and earthwoms (Hagley et al. 1982).

Poecilus lucublandus Say (fonerly known as Pleroaichus lucublandu. Say)

Forbes (1883) obsewed that in the field 114 of P. lucublandus ' food was vegetative. HOW~VWaccording to Best 8 Beegle (1977a) this carabid was reluctant to eat plant material. In the field, the carabid attacked and was observed to feed upon black cutwom lawae (Lund & Turpin 1977), insects (Kirk 1971), E. ochrogaster (Erwin et al. 1976) and earthwoms (Hagley et al. 1982). In laboratory feeding trials P. lucub/andus was observed to feed upon E. ochmgaster, inch woms, moth and buttemy lawae, asparagus beetle eggs (Erwin et al. 1976), adult western corn rootwoms (Diabmtica

M~gifbm)(Kirk 1971), arrot weevils (Baines et al. 1990), second- and fourth-instar annywoms.(Clark et al. 1994). black wtwom larvae (Best & Beegle 1977b), weed seeds (Best & Beegie 1977a), eggs and 5th-instar lawae of E. ochrogaster (Frank 1971a), and live and dead green ctoverwonn lawae (Baines elal. 1990). Lindroth (196 1-1969) ncorded this 9.0-14.0 mm long carabid in open dry areas, however Rivard (1984) found it in moist habitats. It can be found throughout North America (Lindroth 1961 -1989; Kirk 1971)

Pterostichus leconteiunus Lutshnik This 7.9-8.5 mm long carabid has a transarnerican distribution (Lindroth 1961 - 1969). Whereas Lindroth (1961-1969) found it in open dry meadows, Rivard (1964) documented that the beetles occurs in or near woods with moist soil. The feeding habits of the insect are not well recorded.

PterOaicAus luctuosus Dejean Lindroth (1961-1969) reports that this 8-12 mm long carabid occun in moist areas where there is rich vegetation. The beetle has a transarnerican distribution (Lindroth

1961 -1969). The beetle was observed in wptivity to eat mealwoms, a maire kemel a piece of ham and a slug (in Larochelle 1990).

PlelPajchus rndrnarlur llliger Holliday & Hagley (1978) state that P. melanarius is largely camivorous, however some individuels will preferentially eat weed seeds (Hagley et al. 1982). The beetle shows a preference for large prey (Hagley et al. 1982). Under field conditions the. following food items were eaten: grass and grass seed (Johnson & Cameron 1969), arthropodr (Pollet a Desender 7 985) and plant tissue (Pollet & Desender 1985), aphids (Pollet & Desender 1985;Toknen 1995), delphacids (Pollet & Desender 1985; Erwin et el. 1976), earthworms (Hagley et al. 1982; Pollet & Desender 1985), Lepidoptera larvae (Hagley et al. 1982. Pollet 6. Desender 1985), adult tipulids (Pollet & Desender 1985), 0.

146 florallis eggs (Varis1989), and O. radicum (Varis1989). In the taboratory P. melanarius was observed to feed upon grass seed (Erwin et al. 1976), weevils (Hyperodes spp.) larvae. pupae and adults (Erwin et al. 1976). Asparagus beetle eggs (Emvin et al. 1 W6), fruit fly pupae (Tolonen 1995). fifth-instar C. pomonella (Riddick & Mills1994), canot weevil (Baines et al. 1990). weed seeds (Baines et al. 199O), winter moth pupae (Operuphfera brumata) (Frank 1967) and scarab larvae (Hagley et ai. 1982). This 12.û-

19.0 mm long carabid has been recorded in forests (Wallin1986; Lindroth 1961-1969) and on open meadows (Lindroth1961-1969; Rivard 1964), cultivated land (Lindroth 1961- 1969; Rivardl964; Wallin 1986), waste places (Lindroth 1961- 1969). The species was introduced to North America on both Pacific and Atlantic coasts from Europe (Lindroth 196t -1969).

Pterostichus mutus Say Lindroth (1961 -1989) records that the 9.5-1 3.5 mm long P. mutus often occun on dry cultivated soil. It is confined to the eastem half of North America, but has been introduced into southem British Columbia (Lindroth 1961 -1969). Larochelle (1990) notes that in the field the beetle was observed to eat larvae of Leucania unipuneta Haw. (Lepidoptera: Notuidae) and Hetemcampa guttivitta Walk. (Lepidoptera: Notodontidae), and in captivity it was observed to feed upon pieces of ham.

24. SELENOPHORUS Dejean

Sehnophorus elIipticus Dejean Relatively Iittle is known about this 5.36.5 mm long carabid (Lindroth 1961 - 1969). Lindroth (1961-1969) records Mat it is found in dry sandy places. 25. SPHAEROOERUS Dejean

Sphaerodefus hconhl Dejean This 11.5-1 7 mm long carabid is found in eastem North America (Lindroth 1961- 1909). Although it specializes in snail predation (Lindroth A961 -1969), it has been observed to attack cocoons of Swaine jack-pine sawfly (Neopfidion swainec) (Thiele 1977). As well, LarocheMe (1990) lists slugs, mealwoms, earthwoms, ham, banana and apple as some of the food items consurned by the species in captivity. Lindroth (1961- 1969) reports that this is a forest species which prekn moist places with rnosses and dead leaves however according to Rivard (1964) the species occurs in dry soi1 in or near woods.

26. STENOLOPHUS Stephens (Recently placed in genus Agonoderus (Pausch 1979))

Stenolophus comma Fabncius Kirk (1975b) States that this beetle is found in a wide vanety of habitats. The 5.5- 7.5 mm long carabid has a transamerican distribution (Lindroth 1961 -1969). It is omnivorous (Pauch 1979; Hagley et al. 1982). Under laboratory conditions it was observed to feed upon chickweed and dandelion seeds (Hagley et a1.1 Q82),punlane (Hagley et a/.l982), foxtail grass (Hagley et a/.l982), black nightshade (Hagley et a/.1982), 1st-instar codling moth larvae (Hagley et al.l982), apple maggot pupae (Hagley el al.1 SBZ), and early instan of C. pomonella (Riddick 8 Mills1994). In the field S. comma has been knom to damage corn (Riddick & MiIls1994). Steno/ophus conjonctus Say This 3.2-4.3 mm long carabid has a transamerican distribution (Lindroth 1901 - 1969). Lindroth (1961-1909) States the beetle is found in open dry areas with sparse vegetation, however Rivard (1964) records it is found in where there is moderate to dense vegetation. Larochelle (1990) notes that in the field the species was observed to feed upon coccinellids (Coleoptera) and in captivity it was observed to feed upon a piece of ham.. Stenolophus lineo/a Fabriciu s S. lineola is a 7.0-9.1 mm long beetle which has a transameflcan distribution

(Lindroth 1981 -1969). It has been recorded on dry. sandy soi1 (Lindroth 1961 -1969), and has been observed to feed upon insect fragments in the field (Erwin et al. 1976).

Steno/ophus ochmpezus Say S. ochmpezus is found from Manitoba to Nova Scotia, from Utah to Florida (Lindroth 1981 4969). Lindroth (1961-1969) found the beetle in marshes and at the margin of ponds, on son clay or mud, where the vegetation is rich. The 4.8-6.7 mm long species is found south to Flotida, west to Utah north to Manitoba and east to Nova Scotia (Lindroth 1961 -1969). Larochelle (1990) notes that in captivity the beetle was observed to feed upon a kernel of maire. 27. SYNTOMUS (NOTE: FORMERLY KNOWN AS METABLETUS Schmidt-Goebel)

Synfomus amerkanus Dejean Both Lindroth (1961-1969) and Rivard (1964) state this 2.7-3.5 mm long species is found open dry ground with sparse vegetation. The feeding habits of the transamerican carabid are not well recorded.

Tachys lncurvus Say Both Rivard (1964) and Lindroth (1961 -1 969) report that this 2.0-2.5 mm long carabid can be found on open dry ground. The feeding habits of this transamerican beetle are not well recorded.

29. TETRAGONODERUS Dejean

Tetragonodenrr I.scktus Ha Ideman Relatively little has been documented about the ecology of this 4.5-5.0 mm long beetle. Tetmgonoderus Intemectus Gemar The ecology of this 504.0 mm long carabid is not well recorded. 30. TRECHUS Clairville

Tmchus discus Fabricius T. discus was introduced to North America from Europe (Lindroth 1961-1 969). Its distribution in Noroi America was not documented by Lindroth (1961-1969). It has been recorded on slightly moist, usually clayish or peaty ground (Lindroth 1961-1 969). In the field the beetle was observed to feed upon fruit fly pupae (Tolonen 1995).

Tmhus quadristriatus This carabid is widely distributed over al1 of Europe south of the arctic circle, eastwards to the western dope of the Ural mountains (Bousquet et al. 1984). In the south, its distribution reaches North Africa in Morocco and Egypt and continues through Turkey and the Middle East to the Caspian Sea and Tadzhikistan in Soviet Central Asia (Bousquet et al. 1984). In North America, it is found in Ontario and Michigan (Bousquet et al. 1084). The beetle can be found in open fields ( Thiele 1977, Bousquet et a1.1984), as well as many types of forest communities (Bousquet et al. 1984). In the laboratory T. quadristriatus was obsewed to feed upon many types of animal prey but not plant material (Mitchell 1963, Vickennan 8 Sunderland 1975). It feeds on Collembola (Forsythe 1983), mites (Fonythe1983), D. mchcum (Vans 1989), earthwonns (Mitchell 1963) and small beetles (Forsythe 1983). In the field, the beetle was observed to feed upon earthworms (Mitchell 1g63), Collembola (Mitchell 1963; Bousquet et al. 1984). mites (Mitchell 1963; Bousquet et al. ?984),and small beetles (Mitchell 1963; Bousquet et al. 1984), aphids (Bousquet et al. 1984), and eggs and immature stages of cabbage root fly (Bousquet et al. 1984). APPE NDlX 2: KEY TO CAMBIO BEETLE SPECIES COLLECTED FROM SOUTHERN ONTARIO FARM FIELDS

The following working key was developed to help with the identification of the carabid species found in my study (see Chapter 2). Lindroth (1961-1969) and Dillon and Dillon (1961) (the latter for Cicindela spp.) were used to produce the key. Species which are not included in this key because they were identified much later than the rest of the species are: Harpelus faunus, Hapalus funeranus, Agonum funeranus. Agonum piccolum, Agonum cupmum, Amam catinata, Bembidion sordidum, Bembidion tfansversale, and Bembi'dion levettei.

1. Antennae inserted on front of head, huge prominent eyes ...... Cicindela: 2 - Anbnnae inserted laterally ...... 6

2. Elytral punctures contrasting in colour, smallish with dark elytra which have Muelgreen punctures, 11 - 14mm ...... C. punctulata - Punctures not contrasting...... 3

3. Pale markings in anterior half of elytra ...... 4 - Pale markings distally only, bluelgreen metallic elytra with some whitish spots, 10- 16mm...... C. sexgultata 4. Pale rnarkings extended to rnargin, brownish with white markings ...... 5 - Pale markings separate from margin, reddish with white rnarkings along sides of elytra, 12-13mm ...... C. scutella~sIecontei

5. Markings intemipted in apical fourth. 12-13rnm ...... C. mpanda - Markings continuous, large beetle, 16-18mm ...... C. fornosa genemsa

6. Sutural stria recurrent ...... 7 - Sutural stria nomal ...... 9

7. Last segment of maxillary palp reduced, looks similiar to Tmchus quadriJtriatus. 2.0-2.Smm ...... Tachys incurvus - Last segment of maxillary palp normal ...... Trechus: 8

8. Elytra pubescent, 4.4-5.5mm ...... T. discus - Elyta not pubescent, looks very much like Tachys incuivus, but is flatter and more elytral stria are evident, head dark. prothorax dark brown and elytra lighter brown, prothorax slightly denticulate, about 4mm long ...... T. quadristn'atus

9. Elytra tnincate at apex ...... 10 - Elytral apex entire ...... 15

10. Claws pectinete ...... 11 - Claws smooth ...... 13 Il.Elytra and head metallic green, prothorax nanow and brown, tarsi black, 7.0- 8.5mm ...... Calleida puncfata - Elytra pale with dark markings, black prothorax and head, looks like big Bembiidion with broad elytm, 4.5-5.0mrn...... Tefragonoderus: 12

12. Appendages entirely testaceous, between pro-coxae without raised margin ...... T. fasciatus - Between pro-coxae with raised margin ...... T. interseclus

13. Prothorax very long and narrow, elytra with a red-black pattern, 5.8-7.2mm

...... , ...... Co/liu& pensylva ni'ca - Prothorax not long and narrow, rather short, elytra. prothorax and head are very da& ...... 14

14. LOOK SIMILAR - 3rd antennal segment is glaborous (except for apical setae), elytal apex is sinuate, 2.7-3.5mm ...... Syntomus americanus - 3rd antennal segment is pubescent apically, elytral apex is not, or only faintly, sinuate, 3.0-3.4mm ...... Microlestes bmvilobus

15. Last segment of maxillary palp reduced ...... embiio: 16 - Last segment of maxillary palp normal ...... 27

16. Bronzy metallic elytra with distinct .silver-spots'. 4.6-5.6mm ...... 8. ineequele - Elytra otherwise ...... 17 17. Lateral bead of elytra prolonged inside shoulder ...... 18 - Bead pmlonged along base or incomplete at shoulder ...... 20

18. At least outer striae incomplete or missing ...... 19 - EigMh stria evident and complete, 5.0-6.0rnm ...... B. amedcanum

19. Only sutura1 striae complete, strong latero-basal canna, very shiny black, no microsculpture on prothorax and elytra, sometimes, 4.1 -5.3mm ...... 8. nitidum - More than one stria evident and complete, piceous brown, elytra faintly hidescent, 2.8-3.5mm ...... B. obtusum

20. Frontal furrows prolonged on clypeus ...... 21 - Frontal funows not on clypeus ...... 23

21. Base of prothorax straight ...... 22 - Base of prothorax strongly sinuate, small black prothorax and head, brown elytra with 4 dear white spots, 2.8-3.7mm ...... B. quadn~culatum 22. Frontal furrows strong and double on clypeus, larger then B.wnicolor, bblckidaric brown head and prothorax, btown unicolourous elytra, 4.0-4.5mm...... B. oberthuen' - Frontal furows don't reach front of clypeus, small beetle, black prothorax and head, brown elytra with very rough light stripe pattern on dark brown elytra, 2.8- 3.6mm ...... 8. versicolor

23. All dorsal punctures free from striae, metallic fore body 3.84.4mm ...... B. tapidum - Anterior puncture touching third stria ...... 24 24. All stnae complete and evident, 3.7-4.9mm ...... :. ..8. saiebratum - Outer striae incornplete before apex ...... 25

25. Elytra dark with pale spots ...... 26 - Eiytra pale with dark rnarkings. 3.9-5.1mm ...... 6. obscurrsllum

26, LOOK SlMlLAR - Dark head and prothorax, brown elytra with 4 light areas, head lacking microsculpture, 4. f-5.5mm ...... 8. rupicola - Microsculpture evident on base of neck and sides of head, 4.9-6.1 mm ...... B. tetracolum 27. Eleven or more complete striae ...... 28 - Less than eleven striae ...... 31

28. Scutellum concealed by median lobe of prothorax, prosternum very broad, body is almost circular, greenish sheen, black with yellowish rnarkings, 5.4-7.0mrn ...... Omophmn amticanus - Scutellum not concealed, body not alrnost circular...... 29

29. Scutellar stria absent ...... 30 - Scutellar stria present, 19-25mm ...... Calasoma calidum

30. Antennal segments 2-6 strongly setose, black with a greenish or bluish sheen, 7.0- 8.5mm ...... LorFcem pilicomiJ - Labrum deeply emarginate, 11.5-1 7mm...... SphaerPdems lecontei 31. Elytra each with 3 rows of ocellate depression separated by shiny fields ...... Elaphnrs Elytra sculpture otherwise ...... 32

32. Body pedunculate, with fossorial tibia on front legs ...... 33 - Body not pedunculate ...... 38

33. Marginal setae at apex of elytra intemipted, smaller than Clivina...... Dyschin&: 34 - Marginal setae not separated by a gap ...... Clivina fossor

34. One dorsal puncture, black with yellowish antennae, stria obliterated basally, prothorax has greatest width behind the middle, 4.4 -4.9 mm ...... O. erythrocem More than one dorsal puncture ...... 35

35. Two dorsal punctures ...... 36 - Three dorsal punctures, 2.6-3.2mrn ...... D. globulosus

36. Dorsal punctures 1 and 2 present, 3.4-4.2mm ...... O. bmvispinus - Dorsal punctures 2 and 3 present, 3.4-4.9mm ...... D. politus

38. Pale beettes with enlarged pro-tibia1 spun for digging, 13- 17mm ...... Geopinus incmssatus - Pro-tibia1 spurs noml ...... 38

39. Frons mukarinate, eyes prominent, 5.0-5.7mm...... Notiophilus aeneus - Frons otherwise ...... 40 40. Elytra entirely pubescent, metallic ...... 41 - At most outer intervals pubescent (one non-metallic species with broad prothorax entifely pubescent) ...... 43

41. Prothorax nanow, strongly cordate ...... Oxypselaphus pusillus - Prothorax broader...... Chlaenius: 42

42. Head smooth and glabrous, 10.1-13.5mm ...... C. tricdor - Head strongly punctate, 8.0-9.5rnm ...... C. /ithophilus

43. Next to last segment of labial palp bisetose ...... 44 - Next to last segment of labial palp plurisetose ...... 64

44. Claws smooth ...... -45 - Claws serrate, looks like Agonum with long yellowish legs and antennae, elytra are shiny, very sirniliar to C. greganus and C. advena: none of these so far recorded in this collection) ...... Calathus ingrafus

45. Elytral epipleura distinctly crossed ...... 46 - Epipleura not crossed ...... 50

46. Segments 2-3 of antennae carinate ...... Poecilus: 47 - Antennal segments not carinate ...... Ptemstichus: 48 47. LOOKS SlMllAR Prothorax without defined lateral bead, hind angles deplanate, 9.0-14mm ...... P. lucublandus - Lateral bead well defined, hind angles at most nanowly deplanate, prothorax denticulate, 10.5-13.0mm ...... P. chalcites

48. Scutellar stria absent, 7.0-8.5mm ...... P. leconteknus - Scutellar stria present ...... 49

49. Last tarsal segment glabrous underneath ...... 50 - Last tarsal segment with two rows of setae. 12.0-19.0mm...... P. melananus

50. Later-basal fovea of prothorax separated from side margin by a broad convexity, 9.5-t3.5mm ...... P. mutus - Later-basal fovea either extending to side margin or separated from Mis by a narrow carina, 8-12mm ...... P. luctuosus

51 . Frontal furrow prolonged into clypeo-ocular line ...... Steno/ophus: 52 - No clypeo-ocular line ...... 55

52. Setae dorsally on meta-tanal segments 1-4 and ventrally on segment 5, looks like srnall Harpelus with a Mack head, rounded brown shiny pronotum and elytra, abdomen not pubescent, 3.2- 4.3mm ...... S. conjunctus - Meta-tani glabrous where described above ...... 53 53. Tarsi slender,1 st segment of meta-tarsi longer than 2nd. Elytra hes a metallic greenish sheen, head prothorax and elytraare da&, outer edge of pronotum is light, looks like small Harpalus, 4.86.7mm ...... S. ochmpezus - Tarsi short, 1st segment not, or very little longer than 2" ...... 54

54. LOOK SlMllAR Basal bead of prothorax well-defined laterally, usually on the pronotum the dark spots are well separated, 7.0-9.1mm ...... S. lineola - Basal bead not present, 5.5-7.5mm ...... S. comma

55. 2 super orbital punctures, labrum deeply ernarginate, and asymmetrical, black body and prothorax with a well developed lateral bead, 9.7-11.7mm ...... Diplocheils obtusa - Characters not exactly as above ...... 56

56. Mandibles with lateral setae, head with a well delimited constricted head, 9.2-4.8mm ...... Patm bus kngicomis - No mandibular setae ...... Agonum: 57

57. Third antennal segment pubescent ...... 58 - Antennae pubescent from fourth segment ...... 60

58. Elytra iridescent, 8th stria deeply impressed in its Mole length, 4.9-5.9mm ...... A. datiingtoni - Elytra not iridescent, 8th stria shallower esp. in rniddle ...... 59 59. Meta-tarsi with dorsal median furrow, 6.4-9.lrnm ...... A. grefiosum - Meta-tarsi without median fumow, 5.3-6.5 mm ...... A. sordens

60. Upper surface bnghtly metallic, usually with contrasting coloun ...... 61 - Upper surface dark, with at most, a faint bluish/ greenish tinge ...... 62

61. Elytra unicolored, often contrasting with pronotum, individuals Vary widely in colours, 6.4-9.1 mm .... .'...... A. muellen' - Elytral sides contrasting sharply with disc, (red mainly, green sidas) 7.5-9.3mm ...... A. cupnpenne

62. Elytra covered with setigerous punctures, frons with 2 da& rufous spots. Large with long legs ...... A. hypolithus - Elytra not covered with setigerous punctures ...... 63

63. 3 dorsal punctures, faintly metallic, 8.0-9.5mm ...... A. tenue - 4 or more dorsal punctures, not metallic, 6.8-8.8mm ...... A. placidum

64. 2 supra-orbital setiferous punctures ...... Amara: 65 - Supra-orbital pundure single ...... 72

65. Seventh stria with a single apical seta ...... 66 - Seventh stria with at least 2 setae ...... 67 86. Prothorax not as punctate and latero-basal fovea separated from lateral margin by an impunctate convexity or carina, unmetallic. 7.4-lO.8mrn ...... A. latior - Prothorax deeply punctate to lateral margin, fovea not separated by carina, unmetallic, 7.4-9.8mm ...... A. avida

67. Upper surface metallic, terminal segment of al1 palpi black except at tip ...... 68 - Not metallic, palpi not black, 5.2-7.3mm ...... A. rubrica

68. Femora pale, 5.6-7.2mm ...... A. familieris - Femora darker than tibiae ...... 69

69. Basal puncture well separated from lateral bead of prothorax 6.5-9.0mm ...... A. cupreo/ata - Basal puncture close to or touching lateral bead ...... 70

70. Outer basal fovea reduced, inner fovea a sharp line...... 71 - Outer basal fovea deep, linear, 5.3-8.0mm ...... A. conwexa

71. lnner fovea parallel to midline, 6.2-8.8mrn ...... A. aenea - lnner fovea oblique, 6.2-9.3mm ...... A. littodis

72. Second, fifth. and seventh striae with adjacent rows of setae 5.36.5mm ...... Selenophotus ellipticus - Elytra without extra setiferous punctures ...... 73 73. First meta-tarsal segment much longer Man second and third combined, longer than apical $pur...... 74 - Fint meta-tanal segment shorter than or equat to second and third, not longer than apical spur ...... 76

74. Mentum tooth present, appendages pale ...... otiobia: 75 - No mentum tooth, 9.2-10.8mm ...... Anisodacty/us nigenimus

75. Elytra with greenish tinge, contrasüng with pronotum, 8.5-9.1mm ...... N. teminate - Upper surface brownish, not contrasting, 9.0-1 l.Omm ...... N. picea

76. Mentum without tooth, frons with nifous spots, pro-tibial spur angularly dilated basally, frontal fovea linear ...... Anisodactylus: 77 - Protibial spur simple, though often enlarged basally, frons without nifus spots, frontal fovea rounded ...... Harpalus: 78

77. Apical spur of pro-tibia tnfid, 8.8-1 1.Omm ...... A. msticus - Apical spur of pro-tibia simple, 8.3-10.5mm ...... A. sanctaecmis

78. Outer elytral intervals punctate or hairy ...... 79 - Outer intervals smooth and glabrous ...... 84

79. Outer intervals hairy...... 80 - Outer intervals punctate (may be setose also) ...... 81 80. Frons punctate and hairy, 6.5-9.0mm ...... H. puncticeps - Frons glabrous, upper surface metallic ...... H. %finis

81. Basal bead of prothorax complete ...... 82 - Basal bead internipted laterally, 10.1 - 15.2mm ...... H. pennsy/vanicus

82. Punctuation of elytra coven atleast intervals 7-8 evenly. Prothorax constricted towards base...... H. longicoiiis - lntenrals 1-7 either impunctuate, or punctuated at base only, or interval 7 less dense than 8 ...... 83

83. Only base of elytral intervals 2-8 punctate ...... erythropus - Punctuation at base of elytral intervals rather coarse, at least 8. interval punctuate also apically, 10.7-14.2rnm ...... bicoiour

84. No dorsal punctures, very large ...... 85 - Single donal puncture, legs pale, 7.54 1mm ...... H. herbivagus

85. Head densely punctuate laterally behind eyes. Vary large (17.5 to 25mm)...... if. calignosus - Head not punctuate behind eyes 11 to 18 mm long ...... H. enaticus APPENDIX 3: DATA FROM CARABID BEETLE COLLECTIONS

Appendix 3-1. Percent abundance of al1 carabid species collected from high tillage treatment HT1 from 1994 -1 997. The total number of beetles collected was 31 99.

'k abundance Techys incurvus 39.27 Pterostichus melanarius 13.95 Clivine hssor 10.52 Pterostichus lucublandus 8.25 Bembidion quadrimaculatum opposilum 6.64 Agonurn muel/eri 6.53 Bembiûion telracolum 3.32 Stenolophus comm8 2.68 Anisodeclylus sanctaecrucis 1.O8 Trechus quadiistnatus 0.80 Harpalus pensylvankus 0.77 Patmbus longicomis 0.72 L&m pilicomb 0.61 Cicindela sculellsris lecontei 0.55 Fîetuslichus mutus 0.53 Bembidion rspiâum 0.44 Agonurn cupripenne 0.39 Bembidion obtusum 0.36 Geopinus incmssatus 0.30 Hutpalus aRFnis 0.30 Chlaenius tricolor 0.28 Agonum plecidum 0.17 Herpslus hetBivagus 0.17 Synlomus amerkanus 0.17 Bembidion insequele 0.1 1 BembidEon ~mf8 0.1 1 Hatpalus caliginosus 0.t 1 H~W~USPU~C&C)PS 0.1 1 Stenolophus lineole 0.11 Bembidion ve~~icoIor 0.08 TIscnus discus 0.08 Agonum tmue 0.06 Bembidhn am&anum 0.06 Cdliurus psnsyivanria 0.06 Harpslus bngicdlis 0.06 Awum cuprsurn 0.06 Anisodrclylus mstkus 0.03 8etnbiâion nilidum 0.03 8embidion sebbrstum 0.03 Chlmnius lithaphilus 0.03 Omophmn ams~anurn 0.03 Harpalus launus 0.03 Appendix 3-2. Percent abundance of al1 carabid species collected from high Mage treatment HT2 from 1994 -1997. The total number of beetles collected was 7328.

% abundance % abundance Agonum muellen' Agonum darlingtoni Pterostichus melanerius Agonum gratiosum CIivina hssor Agonum hypolithos Bembidion quadrimaculatum oppositum Amers littorelis Loriceta pilicomis Oyschirius brev~spinus Bembidion vemicdor Dyschirius erylhrocerus Bernbidion telmcolum Herpalus bicolor Trechus discus Harpelus caliginosus Pt eroslichus lucublendus Stenolophus lineola Bembidbn obtusum Agonum cupmum Tachys incurwus Amam canhata Stenolophus comma Bembidbn rapidum Agonum cupr@tnne Tfechus qu8drisMatus Harpslus he~Wvagus Anisuûscty/us sanct~~~rucis Bembiâion sabbrsturn Chlaenius Ibbr Hatpalus puncticeps Bsmbidion mprCol8 .. Oyschidus poliius Agonum plscEdurn Bembidion tnnswmuh? Bernbidion oberthueri Aponum tenue Aman N~&U Bsmbidion nitidum Tefregonodemsinfsmbctus Amen cupmdata Anisodactylus nripsmmmus Bernbidkm transwrsale Dyschriius g4bubsus Chleenius Iilhcphilus Haîpelus pensyivanicus Agonum sordbns Aman mma &nela 8w8 Aman /amiliaris hiSOdbc&h~NS~US Cki&le ssxguttata H81pmIusMnis Appendix 3-3. Percent abundance of al1 carabid species collected from medium Mage treatrnent MT from 1994 -1 997. The total number of beelles collected was 3642.

K abundance species % abundance 28.39 Colliunrs psnsylvanica 0.04 14.56 Cicindela punctulata 0.04 9.68 Cicindele mpanda 0.04 7.18 Ciundela sexgutla ta 0.04 7.15 Dyschirius eryîhmcerus 0.04 6.56 Oyschinus bmvispinus 0.04 3.87 Hatpalus puncticeps 0.04 3.75 Harpelus erra ticus 0.04 3.59 Haqaalus bicolor 0.04 1.80 Microlssies bmvilobus 0.04 1.33 Stsnolophus leconlei 0.04 1.21 Herpulus faunus 0.04 1.13 Agonum cuprsum 0.04 0.90 Amam cunneta 0.04 0.86 Bembidion sordidum 0.04 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 O. 12 0.12 0.08 0.08 0.08 0.04 0.04 0.04 0.04 Appendix 3-4. Percent abundance of all carabid species collected from no tillage treatment NT frorn 1994 -1 997. The total number of beetles collected was 6326.

apecimr X abundance species % abundance Pterostichus melanerius 21.95 Harpelus longicollis 0.05 Cicindela punctulata 17.51 Omophron americanum 0.05 Hapalus pensyivanicus 10.28 Petrobus longicornis 0.05 Anisodac~ussanctaecrucis 7.08 Tetragonoderus intersectus 0.05 Bembidion quadnmaculatum oppositum S.? 4 Harpalus faunus 0.03 Clivina fossor 5.31 Amera familiaris 0.03 Plemstichus lucublandus 4.81 Amara littoralis 0.03 Ckindela fonnose generose 4.55 Bembidion nitidum 0.03 Hatpdus hicolor 3.92 Bembidion rapidum 0.03 Cicindele rependa 3.71 Chlaenius ljthophilus 0.03 Harpalus herbivegus 1-96 Geopinus incrassetus 0.03 Ptemstichus luctuosus 1.62 Herpalus erythmpus 0.03 Cicindele scutellens lecontei 1 .O8 Agonum hypolilhos 0.02 Stenolophus comma 0.95 Bernbidion obtusum 0.02 Harpelus afin/$ 0.89 Bembidion versicolor 0.02 Anisudactyrus nrsticus 0.84 Cicindela sexgutfata 0.02 Amara senea 0.70 Colliurus pensylvenica 0.02 Calasoma calidum 0.65 Oyschirius globulosus 0.02 H8fpalus caligiflosus 0.65 Oyschirius obtusa 0.02 Ha~aluspuncticeps . 0.65 Notibia terminala 0.02 Stenolophus canjunctus 0.58 Noliophilus asneus 0.02 Amam mb&a 0.50 Stenolophus lecontei 0.02 Chlsenius In'cdor 0.50 Agonum piccolum 0.02 Bembidion telnicdum 0.48 Amen canna 0.02 Amers au& 0.34 Bembidion sordidum 0.02 Agonum pl~um 0.28 Harpelus funerius 0.02 H~@I~usemtkus 0.21 (VoYbia pics8 0.19 Aman cupmoîata 0.17 LoriCern piIic0mis 0.17 Agonum cupnpenns 0.1 5 Agcwum mueIlen' 0.10 Amam latkw O. 1O Celethus inglstus 0.09 Micmkstes bravilobus 0.09 Re~~~tichusmutus 0.09 Awum sordsns 0.07 Aman conwxa 0.07 Stenobphus /insola 0.07 Awum tenue 0.05 Bembûîian amricanum 0.05 Bambidion inwuab 0.05 L L'9 96'9 L L'O SL'6 a'rr WOt d4.L a's E9'LL U' sz ma: WIL

19'9 W'L 9t'O 80'6 CO'6 L OL -02 SWI

CI'S ws 40'5 40's LW9 WÇC ML