THE OF FLYING COLEOPTERA ASSOCIATED WITH INTEGRATED PEST MANAGEMENT OF THE DOUGLAS-FIR (Dendroctonus pseudotsugae Hopkins) IN INTERIOR DOUGLAS-FIR (Pseudotsuga menziesii Franco).

By

Susanna Lynn Carson

B. Sc., The University of Victoria, 1994

A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

in

THE FACULTY OF GRADUATE STUDIES

(Department of Zoology)

We accept this thesis as conforming

To t(p^-feguired standard

THE UNIVERSITY OF BRITISH COLUMBIA

2002

© Susanna Lynn Carson, 2002 In presenting this thesis in partial fulfilment of the requirements for an advanced

degree at the University of British Columbia, I agree that the Library shall make it

freely available for reference and study. 1 further agree that permission for extensive

copying of this thesis for scholarly purposes may be granted by the head of my

department or by his or her representatives. It is understood that copying or

publication of this thesis for financial gain shall not be allowed without my written

permission.

Department

The University of British Columbia Vancouver, Canada

DE-6 (2/88) Abstract

Increasing forest management resulting from bark beetle attack in British Columbia's forests has created a need to assess the impact of single management on local biodiversity. In the Fort St James Forest District, in central British Columbia, Douglas-fir (Pseudotsuga menziesii Franco) (Fd) grows at the northern limit of its North American range. At the district level the species is rare (representing 1% of timber stands), and in the early 1990's growing populations of the Douglas-fir beetle (Dendroctonus pseudotsuage Hopkins) threatened the loss of all mature Douglas-fir habitat in the district. In response to beetle populations and increasing management needs, forest managers initiated a 5-year operational research study on the impact of pheromone trapping and harvesting on flying beetle diversity.

Beetle diversity was measured from pheromone baited and unbaited Lindgren funnel traps located in mature/old growth, beetle attacked, leading Fd habitat from preharvest, through 4/5th season postharvest conditions. Pheromone traps were baited with known Douglas-fir aggregation pheromones; frontalin, MCOL, and seudenol, and all traps were collected weekly or bimonthly through the duration of the seasonal Douglas-fir beetle flight (April/May through August/September) from 1994-1997.

A total of 484,000 individuals, representing 625 identified species and recognisable taxonomic units (morphospecies), from 67 families were trapped in preharvest and postharvest baited and control sites. Whittaker plots indicate logarithmic species distributions for both pheromone-baited and control trap catches, although the rank position of species varied between treatment conditions and trapping year. Between pheromone-baited and control traps, under preharvest conditions, trap catch analysis resulted in significant differences (cx = 0.05) for eight out of nine

diversity indices including; Margalef's (d), Shannon-Weiner (H'10), Brillouin, Fisher's (rx), Pielou's (J), 1-Simpson's (1-D), Taxonomic diversity (6), and Taxonomic distinctness (8*) . Significant differences between baited and control data across all treatment years (preharvest, post 1, post 2, post 3, post 4/5) were observed for 6 out of 9 indices. Similarities observed in species richness (S) and Margalef's (d) measures are thought to be an artefact of low sampling effort.

Changes observed in diversity and species abundance are thought to have resulted from the disproportionate trapping of an unknown number of non-target species by pheromone-baited traps relative to unbaited traps. Differences in diversity observed across harvest years occurred within the context of, and in addition to, a dynamic and changing species assemblage responding to harvesting and the resulting habitat change. The results suggest that the effect of single species management, in this circumstance, is not limited to the target organism and pheromone trapping along with harvesting as part of an IPM program can influence species composition at the community level. TABLE OF CONTENTS

Abstract ii List of Tables v List of Figures vi Preface / Acknowledgements vii CHAPTER I - Introduction and Overview 1 Sampling Techniques 8 Trap design 8 Sampling Design 10 Trap placement 10 Replication 12 Site Conditions 14 Topography/Climate and weather 14 CHAPTER II - Impact of Aggregation Pheromones on Old Growth Associated Flying Beetle Diversity 17 Introduction 17 Methods 21 Results 29 Discussion 42 Semiochemical composition 56 Trapping efficacy 59 Long term considerations 61 Summary.... 63 CHAPTER III - Impact of Harvesting on Pheromone Biased Diversity Sampling... 65 Introduction 65 Methods 69 Species distribution analysis 74 Diversity analysis 75 Species abundance - baited vs. control 76 Species trends 77 Results 77 Diversity analysis / Overview 100 Species Number (S) 101 Marqalef s (d) 103 Pielou's (J') 104 Brillouin 104 Shannon-Wiener (H'ig) 105 Simpson's M-X) 106 Fisher's (a) 107 Taxonomic Diversity (5) 107 Taxonomic Distinctness (5*) 108 Species abundance-baited vs. control 108 Species trends 110 Discussion 112 Impact of harvesting 113 Surging species abundance 114 Decreasing species abundance 121 Increasing species abundance 123 Depressed species abundance. 124 Harvesting summary 125 Pheromone Effect 128 Natural variation 136 Trapping efficiency 137 Systematic bias 139

iii Semiochemical variation 140 Summary 140 CHAPTER IV - Pheromones and Integrated Pest Management 142 Introduction 142 Pheromones, Insect Ecology, and Community Development 146 Management Impact 148 Containment and mop-up 148 Monitoring 151 Pheromone bias as context 152 Sources of Variation and Experimental Limitations 155 Species variation 155 Habitat variation 157 Study limitations 158 Pheromone composition 159 Site selection and replication 160 Summary 162 BIBIOGRAPHY 166 Appendix I Taxonomic support 183 Appendix II Statistical protocols for Douglas-fir beetle abundance 185 Appendix III Species data by trend 187 Appendix IV Tabular results, Chapter 3 diversity indices 210

iv List of Tables*

Table 1.1 Site replication (preharvest conditions) 14 Table 2.1 List of non-target species known to aggregate to Douglas-fir beetle pheromone components 20 Table 2.2 Site list, trapping year, and biogeoclimatic classification for preharvest pheromone baited and unbaited sites 23 Table 2.3 Summary results of mean abundance/site and total abundance of flying beetle species under preharvest conditions 30 Table 2.M Twenty most abundant flying beetle species (preharvest) 38 Table 2.5 Richness, evenness and dominance measures of flying beetle diversity (preharvest) 41 Table 2M Species of flying Coleoptera statistically more or less abundant betwee pheromone baited and control sites (preharvest) 42 Table 3.1 Site list, trapping year, and biogeoclimatic classification for preharvest and postharvest pheromone baited and unbaited sites 69 Table 3.2 Site replication of baited and control data across treatment years 74 Table 3.3 Site list for 5-replicate diversity analysis, including harvest stage, trapping year, and biogeoclimatic classification 76 Table 3.H Summary results of total abundance and species number of flying (preharvest and postharvest) 79 Table 3.5 Summary results of mean abundance/site and total abundance of flying beetle species (preharvest and postharvest) 80 Table 3.6 Ten most abundant beetle species listed by rank from pheromone baited and control sites 99 Table 3.7 Flying Coleoptera species statistically more or less abundant between pheromone baited and control sites (preharvest and postharvest) 109 Table titles have been abbreviated/paraphrased for length.

v List of Figures*

Figure 1.1 Distribution of Douglas-fir in North America 1 Figure 1.2 Northern Distribution of mature Douglas-fir 2 Figure 1.3 Site Map 13 Figure 2.1 Site Map - preharvest conditions 22 Figure 2.2 Diagrammatic representation of sampling design 25 Figure 2.3 Venn diagram of species distribution (preharvest) 36 Figure 2M Whittaker plot of species abundance (preharvest) 37 Figure 2.5 Whittaker plot of 20 most abundant species (preharvest) 37 Figure 2.6 Results of flying beetle diversity measures (preharvest) 40 Figure 3.1 Mean abundance/site of flying beetles in pheromone baited and control sites (preharvest and postharvest) 96 Figure 3.2 Whittaker plot of species abundance (preharvest and postharvest) 98 Figure 3.3 Results of flying beetle diversity measures in pheromone baited and control sites (preharvest and postharvest) 102 Figure 3.1/ Observed trends in flying Coleoptera trapped in pheromone baited traps under preharvest and postharvest conditions 111 Figure 3.5 Observed trends in flying Coleoptera trapped in unbaited control traps under preharvest and postharvest conditions 111 Figure 3.G Successive phases in the invertebrate community exploiting dead wood 115 Figure 3.7 Phases of ecosystem development after clear-cutting of a second growth northern hardwood forest 126 *Figure titles have been abbreviated/paraphrased for length.

vi Acknowledgements

The project seemed simple enough: Collect some beetle samples, identify them, analyze the results, and write a thesis. Five years, a half a million , 12 taxonomic specialists, 10 field personnel, 15 lab personnel, 7 funding agencies and supporting corporations and 1 baby later, the project is complete. A debt of gratitude goes to everyone who worked on, and supported the task. The logistical, intellectual, financial, and personal support was essential and appreciated.

A special thanks to the efforts of taxonomic specialists from Canada and abroad: Anthony Davis, Ale Smetana, Don Bright, Serge Laplante, Bob Anderson, Ed Becker, Rick Leschen, Fred Andrews, Yves Bousquet, Laurent LeSage, Darren Pollock, and Sean O'keefe. Together these gentlemen represent over 400 years of experience in insect and ecology. Without that expertise this project would have no results. Their knowledge, effort, and interest have made me understand the immense value of human entomological resources. Maintaining their expertise is essential if we are to understand the most basic elements of biodiversity within Canada's natural landscapes.

An additional note of appreciation must be made to project collaborators at The University of Calgary (Dr. Hal Wieser); The University of British Columbia (Dr. Geoff Scudder); and The B. C. Ministry of Forests, Prince George Region (Phil Zacharatos) and Fort St James District (Peter Volk). Their willingness to initiate a project of this magnitude, and support it through to its conclusion, gives me renewed confidence in the future of our forests.

Major funding for this project was provided by: Forest Renewal British Columbia (FRBC); B.C. Ministry of Forests, Fort St James District and Prince George Forest Region; Tanizul Timber; The University of Calgary, The University of British Columbia; and the National Science and Engineering Research Council (NSERC).

-Susanna Guthrie/Carson 15-01-02

vii CHAPTER I Introduction and Overview

Sustainable forest management demands stewardship, and the stewardship of British Columbia's forested lands is rooted in sustaining the productivity and diversity of forest ecosystems. Over the past decade changes in forest management - from single species conservation to sustaining ecosystems at the landscape level - underscore the importance of understanding the impact of management on species diversity. The need to understand management impacts have become increasingly important at the northern range of Douglas-fir (Pseudotsuga menziesii Franco), where epidemic populations of the Douglas-fir beetle (Dendroctonus pseudotsugae Hopkins) have caused habitat loss and increased management in old growth habitat.

Douglas-fir is considered by some

naturalists and scientists to be the most

important forest tree in Western North

America (MacKinnon et al. 1992). The

species has a north-south range of over

4500 Km (2,796 miles) (Figure 1.1).

Extending west to east from the Pacific

coast to the eastern slope of the Rocky

Mountains (Hermann and Lavender

Figure 1.1. Distribution of Douglas- 2001). In the Fort St James Forest fir in North America (•) adapted from Hermann and Lavender (2001).

1 District in central British Columbia, Douglas-fir grows at the northern limit of its North American range. There the Rocky Mountain, Blue, or interior variety (P. menziesii var. glauca (Beissn) Franco), comprises roughly 1% of timber stands in the district, occurring predominantly as mature stands around southern lakes (Figure 1.2).

IDF Biogeoclimatic ecozone

Figure 1.2. Distribution of the northern limit of mature, leading (>f8) Douglas-fir (Pseudotsugae menziesii) in the Fort St James Forest District, British Columbia, along with current Interior Douglas-fir (IDF) biogeoclimatic ecozone designation (•). Inset shows southwest boundary of the Fort St James Forest District, major lakes, and leading Douglas- fir (>f8, >100years old) (•).

Owing to its unique distribution and close proximity to local communities, stands of Douglas-fir in the Fort St James District are highly valued for a range of timber and non-timber values including among others: recreational

2 use, visual quality, biodiversity, and the presence of rare/unique habitats

(Anonymous 1996). A threat to these values came in the early 1990's, when populations of the Douglas-fir beetle grew to epidemic levels in inaccessible northern stands, jeopardizing the survival of all mature

Douglas-fir in the Fort St James District.

The Douglas-fir beetle is the most important bark beetle species associated with Douglas-fir mortality throughout its North American range (Furniss and

Carolin 1977). The beetle is known to successfully breed in felled, injured, or diseased Douglas-fir under endemic conditions, and will utilize apparently healthy trees under epidemic conditions. The beetle also attacks western larch (Larix occidentalis (Nutt.)), but produces brood only in down trees

(Furniss and Carolin 1977). The role of the Douglas-fir beetle in the formation of snags and coarse woody debris (CWD) is that of a primary attacking, phloem boring insect (pioneer saprophage) with secondary attack characteristics (Knight and Heikkenen 1980).

The Douglas-fir beetle is one of 3 Scolytid species (D. pseudotsugae

(Hopkins), Scolytus unispinosus (LeConte), and Pseudohylesinus nebulosus (LeConte)) known to settle on Douglas-fir soon after the death of a tree (Dajoz 2000), The resulting primary attack signals the onset of the first of up to five stages of a tree decay lasting up to 400 years (Maser et al.

1988). Stage one (0 - 6 years) of Douglas-fir decay is characterized by

3 presence of the Douglas-fir beetle and other Coleoptera - predominantly members of the Scolytidae, Cerambycidae, Buprestidae, and occurring in a number of biological relationships including competition, and successional facilitation (Harmon et al. 1986, Berryman 1986, Caza 1993).

Under the bark, the galleries of Douglas-fir beetles are known to be associated with at least 17 beetle species with biological relationships that include; predation on Douglas-fir beetles or other gallery associated insects, fungus feeders, and "unknown relationships" (Deyrup and Gara 1978).

While the list of species associated with beetle attacked, dying Douglas-fir is complex, and in all likelihood not fully understood, perhaps one of the most important species associations of the Douglas-fir beetle in British Columbia is between the beetle and the person responsible for directing the local integrated pest management (IPM) program.

In response to epidemic Douglas-fir beetle populations in the Fort St James forest district, forest managers initiated a district wide, landscape level, integrated pest management (IPM) program that combined harvesting with intensive pheromone baiting/trapping for the Douglas-fir beetle. The primary objective was to reduce beetle populations and retain old growth

Douglas-fir habitat. Secondary objectives included maintaining non-timber values and landscape patterns that would, in turn, maintain multilevel ecological systems associated with old growth Douglas-fir. While forest managers were confident in their knowledge of harvesting, and the impact

4 of harvesting activity on local management priorities, they were less confident in their knowledge of the impact of pheromone baiting.

During the seasonal flight (May through September) of the Douglas-fir beetle's one year life cycle (McMullen and Atkins 1962), the species is known to be strongly influenced by a range of pheromones (Rudinsky et al.

1977). A number of pheromones for the Douglas-fir beetle have been identified to date (Dickens et al. 1985, Ross and Daterman 1995), including three aggregation pheromones (commonly known as MCOL, seudenol and frontalin) which when released in combination produces a synergistic effect on beetle aggregation (Ross and Daterman 1998, Wieser and Dixon 1992) that can be further enhanced with ethanol (Pitman et al.

1975). A survey of separate investigations on MCOL, seudenol and frontalin indicates that one or more of these semiochemicals are associated with 12 flying beetle species (predominantly Scolytidae) with range associations that include , , hemlock, larch, true firs, or Douglas- fir forest habitats (for listing see Table 2.1, Chapter 2).

The natural production of Douglas-fir beetle pheromone components

(including, semiochemical composition, release rates, and geometrical and optical isomers ratios) are thought result from the monoterpene composition of the host plant (Libbey et al. 1985) along with the metabolic processes of the attacking beetle (Grosman etal. 1997). The resulting semiochemical

5 bouquets effect different responses in different beetle species (Gries 1992),

play a critical role in the optimal attraction of conspecifics, and function as a

means of interspecific reproductive isolation (Gries 1992, Paine et al.

1999).

The action of semiochemical systems operating between beetle species within a and populations within a species are considered to be highly

evolved (Wright 1958, Raffa 2001), and will function to mediate host

selection, mate attraction, resource competition, and predation (Hedden et

al. 1976; Chapman 1963; Lanier 1970; Gast et al. 1993; Lessard and

Schmidt 1990; Smith et al. 1990; Rankin and Borden 1991: Herms etal.

1991). In the analysis of reports on pheromone action there is an

awareness of the ecological and economic significance of pheromone use,

but field studies are usually confined to the pheromone-target species

relationship. To date relatively little research has been done on the effects

of pheromones on secondary, non-target species, or on the role of

pheromones in ecosystem development (see Peck et al. 1997). The Fort

St James IPM program created the opportunity to investigate these

questions.

The Fort St James IPM program allowed pheromone researchers to answer

two questions: 1) Do pheromones specific to the Douglas-fir beetle

influence the non-target flying beetle community associated with beetle-

6 attacked mature Douglas-fir, and 2) in the process of managing for epidemic

Douglas-fir beetle populations, is there an impact on non-target flying beetle

species when pheromone trapping is combined with harvesting? Answering these questions involved sampling flying beetles from old growth Douglas-fir

habitat, both in the context of pheromone baiting and in the context of

harvesting.

Assessing the presence and relative abundance of insect species in

response to pheromones is achieved through the use of specially designed

pheromone traps (Muirhead-Thomson 1991), which come in a range of designs and sampling capabilities. All pheromone traps result in the capture

of non-target species though depending on the trap, design can limit sampling efficacy through differences in pheromone dispersal, available

surface area, and behavioural manipulations (Fletchmann etal. 2000). In the context of diversity sampling pheromone traps can be used as a

supplement to traditional trapping methods, or to sample biodiversity using

limited collections of particular taxa (Marshall etal. 1994). A recent study

indicates that pheromone baited Lindgren funnel traps and sticky traps

capture more flying scolytids, cerambycids, and buprestids than window traps (Werner 2002, in press), though the full extent of sampling efficacy

between traditional trapping methods and baited traps has yet to be critically

compared. The specialized nature of the sampling that results from

pheromone trapping, has to date precluded traps and trapping protocols

7 developed for pheromone use from the guidelines of the British Columbia

Resources Inventory Committee (RIC) (Winchester and Scudder 1993).

Pheromone trapping comes with its own unique requirements including trap designs, sampling protocols and sampling limitations. In order to provide context for the type of non-conventional, multi-species, multi-habitat assessment proposed in the Fort St James IPM study, it was necessary to integrate pheromone requirements with the essential elements of field sampling. Elements for integration include: sampling techniques (trap design); sampling design (trap placement and replication); and site conditions (climate and topography).

Sampling Techniques

Trap design

Studies indicate that changes in community structure can be assessed through easily sampled insect communities (Lehmkuhl etal. 1984). Such multi-species assessments are traditionally accomplished through a range of passive flight interception traps (Winchester and Scudder 1993) such as window flight traps (Chapman and Kinghorn 1955) or Malaise traps

(Hutcheson 1990). Studies indicate that the spectrum of captured is sensitive to the choice of trapping method (Trueman and

Cranston 1997) and within trapping systems the highest entry occurs in traps with a large surface area for interception and omni-directional access

8 (Muirhead-Thomson 1991). Compared to traditional methods for surveying

flying arthropods such as Malaise traps and window traps, pheromone traps

have a smaller catch surface area, reducing the potential for random flight

interception and sampling. For species of interest, pheromone trapping

creates differential attraction of species responding to a pheromone plume that is known to increase sampling efficiency by increasing the effective trap

radius, but this effect is variable and present only for those species

exhibiting a pheromone response (Byers etal. 1989). For assessing

pheromone use beyond the target species, sampling requires a trap able to

disperse pheromones and sample flying beetle species with minimal

maintenance requirements. Within the range of available pheromone traps,

the Lindgren multiple funnel trap is an effective trap to satisfy sampling

requirements.

While the dispersal of pheromones in field conditions is ultimately governed

by natural physical and meteorological conditions (Elkinton and Carde

1984), Lindgren funnel traps have been shown to provide optimal chemical

dispersion regardless of lateral wind direction (Lindgren 1983). The traps

are low maintenance, and are known to sample both target and non-target

species (McLean et al. 1987, Ross and Daterman 1998), predominantly

Coleoptera (Lindgren 1983). Studies indicate that trap coloration can have

a positive or negative visual influence on different species/families of flying

insects (Kirk 1984), and black traps (the colour of multiple-funnel traps) are

9 an appropriate colour for target and non-target beetle sampling (Dubbel et al. 1985). Comparing pheromone baited, Lindgren multiple-funnel

pheromone traps to pheromone baited slot traps for Douglas-fir beetles

resulted in a significantly greater number of Douglas-fir beetles and non- target predators in multiple-funnel traps as measured by total numbers/trap

(Ross and Daterman 1998). This result is consistent with comparative studies of pheromone traps in other forest types (Fletchmann et al. 2000), and the increase in target and non-target sampling from individual multiple- funnel traps is thought to result from a greater total surface area/trap than other trap designs, effectively increasing the traps potential for both active and passive flight interception.

Sampling design

Trap placement

Trapping for flying Coleoptera is a three-dimensional problem that requires

optimum vertical and horizontal trap placement. In forest environments the

distribution of insects occurs along a vertical gradient that changes with

species and habitat parameters including forest structure. In stands with

variable stand retention (including clear cut conditions), insect richness was

observed to decrease with increasing trap height above 1 meter (Su and

Woods 2001). Optimum adult trapping of ambrosia beetles (Trypodendron

linneatum (Olivier)) is known to occur at or just below the height of adjacent

underbrush, from 1-2.5m (Shore and McLean 1984) underscoring the

10 influence of forest structure on insect movement. The distribution of flying scolytids in oak- forest found the highest species richness occurred

1 -2m off of the forest floor (Roling and Kearby 1975). These findings are consistent with optimum 1.5m trap height reported for pine beetle (Tilden et al. 1979), and initial attack height the of 1-2m observed for Douglas-fir beetles (Prenzel et al. 1999).

Unlike vertical placement, horizontal trap placement across a forested

landscape considered two scales of measurement; 1) within site trap distance, and 2) between site distance. Within site trap distance was set at a minimum of 50m based on known orientation distances, and aggregation

radii of various Scolytid beetles in response to pheromones. The spruce beetle (Dendroctonus rufipennis (Kirby)) is known to orient to conspecific pheromones from up to 17m (Byers 1995), and the species has an effective

25m range of semiochemical attraction (Shore er al. 1990). These

numbers are consistent with an observed Douglas-fir beetle spillover attack

radii of 20.1m and 22m surrounding pheromone baited trees (Ringold et al.

1975, Thier and Weatherby 1991). The ability of baited traps to induce a differential attraction of species in response to aggregation pheromones is thought to increase sampling efficiency by increasing the effective trap

radius (Byers et al. 1989), so by setting distances based on the responses

of the target species (presumably the most influenced species), any

pheromone influence on non-target species should be less than the range of

11 influence for the target species both within and between sites. Between site distances ranged from 150m to 30km, and were determined by the location

and accessibility of harvestable, beetle attacked stands. All available sites

in the district that met requirements for trap replication, spacing, and sampling frequency were included in the study (Figure 1.3).

Replication

Replication is one factor not directly affected by the influence of

pheromones. However, the replication applied to this study is worth

addressing because it has no precedent in the scientific literature. The study gathered data from more than 270 traps in 67 sites representing 5 treatment years/conditions (pre-harvest, 1st season post-harvest, 2nd season post-harvest, 3rd season post-harvest and 4/5th season post-harvest

conditions). Data were gathered over the duration of the main flight period

of the Douglas-fir beetle (May through August/September, depending on

seasonal flight variations) from a minimum of four traps per site, from which three traps would be used for data analysis. Samples gathered from the three traps were combined to generate a single data set for each site.

Utilizing a combination of three traps/site was based on maximizing species

representation for each site with limited resources, while eliminating

concerns of pseudoreplication. Over-sampling of sites at the beginning was

initiated because of the expected one in four loss of trap samples owing to

poor trap construction, or damage.

12 CP 32/80 (1) CP 120/2 (3) CP 46/222 (3) Kuz Che (1) CP 123 (3) CP 118/1 (3) CP 115 (6) Hobson Is. (5) TachieHill(5) Germajisen-Pinchi (10) PinchiHill(2) Tachie-Pinchi (5) RNW (2) RNE (2) AP1-4,APC,CP18 (10) Whitefish(5) Siesmic (1)

Figure 1.3. Location of 17 study areas, containing 67 sites, initiated between 1994 and 1997 in the Fort St James Forest District, British Columbia. The number of sites per study area indicated by ().

As mentioned above, the study gathered data from 67 sites representing 5 treatment years/conditions. Site replication ranged from 5 to 11 sites per treatment (Table1.1). Aware that unequal replication strongly influences sampling intensity and diversity calculations (Magurran 1988), the study assessed equal replication 'subsets' of the data for all affected analyses.

Assessing questions of pheromone impact, alone and in conjunction with harvesting, is achieved through the use of two data sets: one data set limited to preharvest analysis with higher replication, and a second data set that combines preharvest and post harvest analysis with lower replication.

13 Differences in replication mean that the two analyses (presented in two separate chapters), although related, are not directly comparable. To emphasize the self contained nature of each data set, full details of the study methodology (including replication) are presented separately for each chapter despite the inevitable redundancies.

Table 1.1. Site replication of baited and control data across treatment years for the Fort St James pheromone study. * Indicates sites in which diversity calculations are included for general comparison only. Treatment Preharvest Post 1 Post 2 Post 3 Post 4/5 year Baited or bait control bait control bait control bait control bait control control # of site replicates 10 7 11 1* 11 1* 10 5 6 5

Site Conditions

Topography/Climate & weather

Because of the nature of pheromones as airborne volatiles, one of the primary factors affecting trapping efficacy is the presence, intensity and duration of climatic factors, particularly temperature and wind (Farkas and

Shorey 1974). In a field study environment neither of these factors can be controlled. However their potential influence makes it prudent to assess for seasonal variations, regional extremes or potential anomalies.

Seasonal climate variation is perhaps the most unpredictable factor affecting beetle emergence, activity and pheromone efficacy in natural

14 environments. To accommodate seasonal variation in the Fort St James study, field trials were carried out over four consecutive seasons from 1994 through 1997. Each season was subsequently assessed for extended

(>4days) extreme weather conditions (daytime (13:00) temperature >30°c or

< -5°c) that might unduly bias study data. None were found.

The potential influence of wind on beetle movement was also considered.

Geographical/topographical features are the dominant influence on local wind patterns affecting insect migration (Burt and Pedgley 1997). Broad shallow valleys containing large lakes characterize the topography of the

Fort St James District. Local convective circulations associated with lakes and hillside drainage currents are thought to be the most likely vector of short-range dispersal. Seasonal prevailing, "over flow" winds are thought to be the dominant medium to long range dispersal influence. The predominance of lakes adjacent to stands of Douglas-fir, and the lack of topographic extremes suggests similar wind patterns for study sites throughout the district. Although individual species distributions and dispersal patterns are not known and cannot be estimated, the lack of extremes suggest the potential for sampling species within a habitat, across the district is not thought to be subject to extreme bias by wind dispersal.

The large body of research available for Douglas-fir beetles, their

pheromone systems, and the physics of dispersal made it possible to

15 integrate pheromone and non-pheromone sampling protocols. Sampling techniques, sampling design, and site conditions accommodate the essential parameters of both trapping regimes, allowing for the development of a protocol to assess non-target trap catches associated with pheromone

baiting for Douglas-fir beetle across a range of Douglas-fir beetle associated

habitats. Assessing the non-target flying beetle community associated with

beetle attacked mature Douglas-fir - preharvest conditions - (Chapter 2), and in the context of harvesting of old growth stands through the initial stage of Douglas-fir decomposition - preharvest through 4/5th season postharvest conditions - (Chapter 3).

16 CHAPTER II Impact of Aggregation Pheromones on Old Growth Associated Flying Beetle Diversity

Introduction

Within insect populations, pheromone reception is known to control and/or

modify behaviour. Of all the behavioural responses elicited by pheromones,

the ability to attract a desired species has become an important tool for

monitoring and manipulating species known to impact human resources. The

Douglas-fir beetle (Dendroctonus pseudotsugae) is the most damaging beetle

species to mature Douglas-fir (Pseudotsuga menziesii) in North America. The

beetle's ability to cause tree mortality, and the resulting economic impact has

made the species' life history and ecology an area of research since the

1950's (Rudinsky 1966b). Since 1965, independent investigations have

identified eight aggregative or anti-aggregative pheromones generated by female/male Douglas-fir beetles (Ross and Daterman 1995). Laboratory and field studies have proven three of the eight identified pheromones: MCOL (1-

methylcyclohex-2-en-1-ol) (Libbey et al. 1983), seudenol (3-methylcyclohex-

2-en-1-ol) (Dickens etal. 1984) and frontalin (1,5,-dimethyl-6,8-

dioxabicyclo[3.2.1]octane) (Pitman and Vite 1970), to be highly effective in

aggregating Douglas-fir beetles (Prenzel et al. 1999; Ross and Daterman

1998)

Aggregation pheromones play a critical role in the lifecycle of the Douglas-fir

beetle. Airborne chemicals released by actively burrowing females mitigate

17 and regulate host selection, mate attraction, and intraspecific competition over the insect's one-year life cycle (Dickens et al. 1984). Every spring adult

Douglas-fir beetles emerge from a temperature-mediated hibernation

(McMullen and Atkins 1962). Pioneer females disperse before locating potentially good breeding habitat through the reception of tree kairomones

(Heikkenen and Hrutfiord 1965). As the female successfully excavates a brood gallery in the sapwood, her frass releases a chemical bouquet (Kinzer etal. 1971), signaling to conspecifics the location of suitable habitat for brood production, and to potential mates, the location of a suitable female (Rudinsky et al. 1977). During the colonisation process pheromone release is modified to include both aggregants and antiaggregants (Pitman and Vite 1974), creating a dynamic, density dependent, self regulating system of resource utilization. Field applications of synthetic aggregation pheromones are known to influence Douglas-fir beetle attack patterns in standing timber throughout their seasonal flight period (Ross and Daterman 1997; Thier and Whetherby

1991), beginning in the spring (April - June) and ending in late summer

(August - September), depending on local and seasonal climate patterns

(McMullen and Atkins 1962, Prenzel etal. 1999; Ross and Daterman 1997;

Lessard and Schmid 1990).

When tested individually, the semiochemicals of the Douglas-fir beetle lure

(frontalin, MCOL and seudenol) are known to elicit behavioural responses from a number of scoytid beetles (Table 2.1). Under field conditions the

18 simultaneous release of frontalin, MCOL and seudenol, as a ternary lure or in combination with ethanol has shown strong efficacy for initiating an aggregation response in populations of Douglas-fir beetles in southeastern

British Columbia, north-central British Columbia, and north-eastern Oregon

(Prenzel etal. 1999; Guthrie and Wieser 1994; Ross and Daterman 1995,

respectively).

When field testing pheromone efficacy, secondary (i.e. non-target) species are

often found in addition to the target species (Wieser and Dixon 1992;

Zahradnik 1995; Peck et al. 1997), though most controlled studies on

pheromone action confine their context to the pheromone - target species

relationship. Despite an awareness of the ecological significance of

pheromone use (Vite and Baader 1990), and studies citing significant, and

disproportionate effects of bark beetle pheromone lures on clerid predators

(Furniss et al. 1974, Ross and Daterman 1995), little research has been

done to elucidate the full extent of pheromone action on non-target species

beyond known predators, or other scolytid beetles.

In 1993, a four year pheromone research project was initiated in the Fort St

James Forest District, Fort St James, British Columbia. One of the objectives

of the project was to assess the impact of Douglas-fir beetle pheromones on

the species diversity of flying Coleoptera associated with beetle attacked,

mature Interior Douglas-fir habitat. The null hypothesis was that pheromone

19 baiting would have no impact on the beetle community beyond the target species.

Table 2.1. List of non-target beetle species known to aggregate to Douglas-fir volatiles (including ethanol), Douglas-fir beetle attacked mature Douglas-fir habitat, and known Douglas-fir beetle aggregation pheromones (MCOL, Frontalin & Seudenol). Listed tree species indicate associated study habitat / known habitat limitation.

Identified Source of Attraction Beetle Douglas- attacked fir Douglas- volatile Seud• fir +EtOH MCOL Fn enol References

1 2 3 4 5 1-5 Species Dendroctonus u,J,4Wood 1982 pseudotsugae 4 Dickens ef al. 1985, Hopkins Lindgren 1992. # 5 (Scolytidae) • • • • Pitman etal. 1975, Ross and Daterman 1995. Dendroctonus 4 Wood 1982. brevicomis Lec. » (Scolytidae) Dendroctonus 4 Wood 1982, Payne et frontalis Zimm. • al. 1978, Payne etal. Pine (Scolytidae) 1988. Dendroctonus J Borden etal. 1990 ponderosae • (contrary to Libbey et Hopkins Pine al. 1985) (Scolytidae) Dendroctonus J Borden etal. 1996 rufipennis Kirby 4 Dyer 1973, Lindgren (Scolytidae) • • • 1992. Spruce Spruce Spruce 5 Furniss etal. 1976. 3,4,5 Setter and Borden 1999. Dendroctonus # 4 Payne etal. 1987, terebrans (Olivier) Delorme and Payne Pine (Scolytidae) 1990. Hylastes nigrinus 'Rudinsky & Zethner- Mann. • • Moller 1967. (Scolytidae) 2Rudinsky 1966a. Hylates ruber ^Rudinsky 1966a. Swaine • (Scolytidae) Trypodendron z Rudinsky 1966a. Lineatum Oliv. • 4 Lindgren 1992, Setter (Scolytidae) • and Borden 1992. Dryocetes ^Rudinsky 1966a. autographus Sw. m (Scolytidae) Gnathotrichus • ^Rudinsky 1966a. 20 retusus Lec. (Scolytidae) 9 Gnathotrichus ^Rudinsky 1966a. sulcatus Lec. • (Scolytidae) Pseudohylesinus 'Rudinsky 1966a. grandis Sw. • (Scolytidae) Pseudohylesinus z Furniss et al. 1974, nebulosus LeC. Rudinsky 1966a. (Scolytidae) dubius 4 Wood 1982, Dixon & LeC. • Payne 1980. Pine () Thanasimus ' Furniss etal. 1974. undatulus Say. • 4 Wood 1982, (Cleridae) Lindgren 1992. Enoclerus 1 Furniss ef al. 1974. sphegeus Fab. • (Cleridae) Abraeus sp # 4 Dixon & Payne 1980. () Pine Cylistix attenuata • 4 Dixon & Payne 1980. (Histeridae) Pine Platypus * Dixon & Payne 1980. flavicomis Herbst • Pine (Platypodidae)

Methods

Samples of flying beetles were gathered from fourteen preharvest sites in the

Fort St James Forest District, Fort St James, British Columbia, between 1993

and 1997 (Figure 2.1): Site replication included seven site replicates for

pheromone-baited traps and seven site replicates for unbaited, control traps.

Two of seven replicates (baited and control sites) were a matched pair for trapping year, site location and habitat. Three replicates were matched for

site location and habitat in different years, and two replicates were matched

by biogeoclimatic designation. The sites chosen included all accessible and

harvestable Douglas-fir beetle attacked stands in the district.

21 CP 120/2 (2)

Kuz Che (1)

CP 118/1 (2) CP 115(1)

Germansen-Pinchi (2)

Tachie-Pinchi (2)

CP 18 (1) Whitefish (2) Siesmic (1)

Figure2.1. Location of preharvest study areas, initiated between 1994 and 1997 in the Fort

St James Forest District, British Columbia. Dots indicate all majority (F8 or greater), mature to over mature Douglas-fir in the Fort St James Forest District. Solid shading indicates location of major water bodies. Left boundary line indicates western limit of forest district. The number of sites per study area are indicated by ().

At all sites, 12 funnel Lindgren traps (Phero Tech Inc., Delta, B.C.) were used for both pheromone baited and unbaited (control) flight interception. Trap protocol required that a standard minimum of four traps per site be placed at least 50m apart and at least 50m inside the habitat margin. Traps were deployed so that collection cups were suspended 1 -1.5 m above the ground, clear from interference by understory plants (Figure 2.2). All collection cups contained a 3cm2 piece of neuro-insecticide impregnated plastic (Vapona brand, dichlorvos (2,2-dichclorovinyl dimethyl phosphate)) to prevent insect escape, and reduce necrophagous activity. Collection cups released

22 rainwater through a bottom screen to create a dry trapping system. No

British Columbia Resource Inventory Committee (RIC) standards are available for this sampling technique.

Baited and control traps were placed within leading or pure (greater than 80%)

Douglas-fir stands (>Fd8), in the biogeoclimatic subzones SBS dw, SBSdk,

SBS wk, & SBSmk (Meidinger and Pojoar 1991) (Table 2.2). Stand age was mature to over mature (110 - 350 years) and stands had not been previously harvested. The trapping season covered the flight season of the Douglas-fir beetle. Sites were initiated in late April/early May, and were maintained until mid to late August. The timing of trap placement was determined by climate and site factors including snow pack and road conditions. Despite yearly variation, all site sampling was implemented prior to the onset of the main

Douglas-fir beetle flight. Traps were removed after both flight peaks of the

Douglas-fir beetle had passed and field personnel observed 2-3 weeks of low

Table 2.2. Site list, trapping year, and biogeoclimatic classification for preharvest pheromone baited and unbaited sites in the Fort St James Forest District, British Columbia Site Number Site Name/Location Sampling Year Biogeoclimatic sub zone 1 Tachi-Pinchi 1994 SBSdw3 1c Tachi-Pinchi 1997 SBSdw3 2 Germansen-Pinchi 9 km 1994 SBSmk 2c Germansen-Pinchi 8 km 1997 SBSmk 3 TFL CP 115 1996 SBSwk3 4 TFL CP 120 1997 SBSdw3 4c TFL CP 120 1997 SBSdw3 5 TFL CP 118 1996 SBSwk3 5c TFL CP 118 1997 SBSwk3 6 Kuz Che 1996 SBSwk3 7 Whitefish D 1997 SBSwk3 7c Whitefish D 1997 SBSwk3 8c Apollo CP 18 1997 SBSwk01/04 9c Seismic Trail 1997 SBSdk

23 to no Douglas-fir beetle numbers at all sites. Traps were emptied weekly,

bimonthly, or monthly as determined by schedule or site accessibility.

Pheromone traps were baited with a Douglas-fir beetle aggregation lure developed by researchers at the University of Calgary. The lure consists of a ternary blend of racemic (±) frontalin (Fn), racemic (*) MCOL and seudenol of an undetermined enantiomeric composition. The release rate of frontalin was

independently regulated from the release rate of MCOL and seudenol. The

release rate of1 Fn was 0.3 mg/day from capillary tubes of 1.0 mm diameter.

The MCOL-seudenol blend was achieved by dispensing pure - MCOL

released at an average rate of 3.0 mg/day from a microcenterfuge tube with a

2mm hole in the cap. The resulting open system of dispersal allowed MCOL to react with atmospheric water (thought to result from condensation) to

produce an MCOL-seudenol interconversion of an undetermined rate that was

observed to stabilize at a 40-60 ratio (respectively). Chemicals were placed

in a hooded cradle attached inside the third lowest funnel of the Lindgren trap

(Figure 2.2). Lures contained enough semiochemical for the duration of the trapping season and were only changed in response to animal damage, or

due to random selection for gas chromatography analysis (to monitor

chemical integrity).

24 Figure 2.2. Diagrammatic representation of sampling design for the assessment of flying beetle biodiversity in mature interior Douglas-fir. Far right image shows forest cover including stand composition, road access, and cut block location. The four stars represent individual Lindgren 12-funnel traps placed at intervals of 50m with collection cups 1-1.5m off of the forest floor. Left image shows cradle design and pheromone placement in the trap.

Following collection, samples were stored in heavy-duty "Ziploc" freezer bags, and frozen to minimize desiccation. Samples remained in frozen storage until shipment to The University of Calgary for the first of two sorting procedures.

Following frozen storage, samples were washed and disinfected in 70% ethanol for a minimum of 30 minutes and strained using a 1mm wire mesh.

Samples were then dried at room temperature in a fume hood from one to four hours (time dependent on sample size), and hand sorted with the aid of a dissecting microscope. This initial sort separated out the target species (the

Douglas-fir beetle) from other Coleoptera and removed obvious debris from the samples. Target species abundance was estimated by weight through regression analysis (Appendix II) and target species identification was

25 achieved through either census or sampling depending on the number of beetles contained within the sample. For samples with 100 or less

Dendroctonus beetles, identifications were achieved through census, while identification of samples with greater than 100 Dendroctonus beetles was estimated through subsampling. Samples containing greater than 100

Dendroctonus specimens were themselves sampled for species composition by identifying the first 100 Dendroctonus beetles sorted from the sample.

Non-target species were re-frozen and transported to the University of British

Columbia for final sorting, mounting, and identification.

Beetle identifications were accomplished by the use of available keys, and by reference to named specimens in both the Spencer Entomological Museum

(University of British Columbia) and the Canadian National Collection of

Insects (Agriculture and Agri-Food Canada, Ottawa). Species and species groups/recognizable taxonomic units (RTU) were identified through the assistance of taxonomic specialists in Canada, the United States, and New

Zealand. Coleopterists at Agriculture and Agri-Food Canada in Ottawa identified specimens for the majority of species, creating a voucher collection for subsequent identification. A listing of specialists and their assistance to this project is contained in Appendix I.

Following identification of all specimens, collection and sample data were entered into a spreadsheet. Data were reduced to a standard of three traps

26 per site. Trap selection was determined either by missing sample (eg, broken/damaged trap), or by random deletion of existing data. Data editing was limited to the removal of necrophagous species suspected to be a direct artifact of the trapping process, or species whos abundance might have been altered as the result of processing protocols (such as the inclusion of species smaller than the 1mm mesh size used in sample cleaning). Editing was

limited to these criterion because the objective of the study was to assess all

non-target species potentially influenced by pheromone trapping regardless of the nature of the association. Data analysis then assessed rank abundance, diversity at the community level (including richness, evenness and dominance), and abundance at the species level.

A total of 9 measures of diversity were applied to the data to assess richness,

evenness, dominance and taxonomic diversity. The use of multiple indices was determined to be appropriate for reasons of uncertainty over the impact

of dominant vs. rare species in sample collections, concerns over the effect of

sample size, and the desire to test recently developed indices based on

taxonomic criteria. Measures of richness and diversity were calculated with

the software program - PRIMER (Plymouth Marine laboratory, Plymouth, UK).

With no previous literature available on the sampling capability of the Lindgren

funnel trap, or on the diversity of the flying beetle community of northern

interior Douglas-fir, existing data to determine minimum sample size was

27 unavailable. The sampling intensity of preharvest sites was moderately high based on the number of traps (21 each of baited and control), but data analysis was pooled by site to reduce concerns of pseudoreplication, effectively reducing sample size to 7 sites in each treatment. The resulting uncertainty over sampling effort was accommodated for with the selection of multiple indices based on their sensitivity to sample size (Magurran 1988).

For richness indices, S (species richness) is considered to be the most sensitive to sample size as it directly reflects the sampling curve. Margalef s index is also highly sensitive to sample size but its calculation is less influenced by rare species. The Shannon and Brillouin indices have moderate sensitivity, while Fisher's

"taxonomic relatedness" (based on a measure of taxonomic distance within species assemblages) into their calculations.

To address the requirement of equal sample size for comparing diversity indices, results presented at the community level were generated with an equal replicate subset (7, 7; baited, control replicates) of the larger, complete data set (10, 7; baited, control). Comparisons of baited and control indices

28 were then subject to statistical analysis by a T-test for difference = 0 (vs not =

0) using the software program MINITAB 2002. Assessment at the species

level included a non-parametric analysis (Wilcoxon Rank-sum test) of

abundance for baited and control traps. Species-level analysis consisted of the entire data set.

Results

A total of 99,467 individuals from 241 species + 29 recognizable taxonomic

units (RTU) or morphospecies, representing 47 families, were trapped in ten

baited and seven unbaited sites. Out of the total species complement, three

species {Nicrophorus diffodiens Mannerheim, Nicrophorus investigator

Zetterstedt, and Catops egenus (Horn)) representing 3461 individuals were

identified as necrophagous in nature and were removed from the data. An

additional eight species (16 individuals) with a body size <1mm (C/'s striolatus

Casey, Cryptophorus sp., Dolichecis indistinctus Hatch (); Atomaria sp.

1,5,8, (Cryptophagidae); Corticaria gibbosa (Herbst), and Lathridius hirtus

Gyllenhal (Lathridiidae) were removed because of sorting inefficiencies,

leaving 95,990 individuals from 259 species/RTUs for analysis (Table 2.3).

The equal replicate subsets of seven baited and seven control sites (used for

all analyses except non-parametric Wilcoxon Rank-Sum analysis) contained

72,546 individuals from 249 species/RTUs.

29 Table 2.3. Summary results of mean abundance per site and total abundance of flyng beetles trapped in baited and unbaited Lindgren funnel traps in mature Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained Mcol, seudenol, & frontalin pheromones in a Douglas-fir beetle (Dendroctonus pseudotsugae) aggregation lure.

Mean Mean Total # # number per per caught site site Baited + Family Genus species Baited Control Control Anobiidae Caenocara scymnoides LeConte 0.2 0.0 2 Dorcatoma (prob) americana 0.4 0.1 5 Hemicoelus carinatus (Say) 0 0.1 1 Microbregma e. emarginatum (Duftschmid) 0.6 0.7 11 Utobium elegans (Horn) 0.1 0.0 1 Buprestidae Anthaxia inornata (Randall) 0.2 0.1 3 Byrrhidae Curimopsis sp. 0.1 0.0 1 Bhyrrhus sp. 0 0.1 1 Cantharidae Podabrus piniphilus (Eschscholtz) 0.1 0.6 5 Silis d. difficilis LeConte 0.1 0.0 1 Carabidae Amara idahoana (Casey) 0.1 0.0 1 nigrinus (Dejean) 0.1 0.0 1 advena (Leconte) 0.1 0.1 2 Pterostichus adstrictus Eschscholtz 0 0.1 1 Sericoda quadripunctata (DeGeer) 0.1 0.0 1 Syntomus americanus (Dejean) 0.1 0.0 1 Cephaloidae tenuicorne LeConte 0.3 0.6 7 Cerambycidae Asemum striatum (Linneaus) 0.1 0.1 2 Corcodera (prob) longicornis (Kirby) 0 0.1 1 Cortodera m. militaris (LeConte) 0.4 0.0 4 Dicentrus bluthneri LeConte 0.2 0.3 4 Evodinus monticola vancouveri Casey 2.7 0.9 33 Grammoptera subargentata (Kirby) 0 0.1 1 Judolia m. montivagens (Couper) 0 0.1 1 Megasemum asperum (LeConte) 0.1 0.6 5 Neanthophlax mirificus (Bland) 0.3 1.7 15 Pidonia scripta (LeConte) 0.5 0.1 6 Phymatodes dimidiatus (Kirby) 0.5 0.1 6 Phymatodes maculicollis LeConte 0 0.1 1 Pygoleptura n. nigrella (Say) 0.1 0.0 1 Spondylis upiformis Mannerheim 4.3 3.3 66 Strictoieptura canadensis cribripennis (LeConteJ 0.1 0.0 1

30 (leConteJ velutinum LeConte 0.4 1.3 13 Trachysida a. aspera (LeConte) 0.3 0.3 5 Xylotrechus longitarsis/undatulus (Casey/Say) 0 0.1 1 Cerylonidae Cerylon castaneum Say 0.6 0.9 12 Chrysomelidae Orsodacne sp. 0 0.1 1 Orsodacne atra (Ahrens) 0.1 0.6 5 Syneta pilosa W.J. Brown 0 0.3 2 Syneta albida LeConte 0.3 0.6 7 Syneta hamata Horn 0 0.1 1 Ciidae Cis sp. (fuscipes) Mellie 0.3 0.0 3 Dolichocis manitoba Dury 0.1 0.0 1 Orthocis punctatus Casey 1.4 0.3 16 Cleridae Enoclerus sphegeus (Fabricus) 0.2 0.0 2 Enoclerus nr. scheaferi Barr 0.1 0.0 1 Thanasimus undatulus (Say) 120 2.0 1216 Coccinellidae Mulsantina picta (Randall) 0.2 0.0 2 Psyllobora vigintimaculata (Say) 0 0.1 1 Colydiidae Lasconotus intncatus Kraus 0.1 0.0 1 Corylophidae Molamba obesa Casey 0 0.1 1 Saeium lugubre LeConte 0 0.1 1 Cryptophagidae Antherophagus sp. # 1 0.4 0.0 4 Antherophagus sp. # 2 0.1 0.0 1 Atomana sp. # 12 0.1 0.0 1 Caenocelis sp. # 1 0.3 0.3 5 Cryptophagus sp. # 1 0.4 0.3 6 Cryptophagus sp. # 2 0.4 0.0 4 Cryptophagus sp. # 3 0.3 0.7 8 Cryptophagus sp. if 4 0.7 2.1 22 Henoticus sp. if 1 0.1 0.0 1 Salebius nr. Minax 0.4 0.6 8 Cucujus claviceps Mannerheim 4 1.9 53 Dendrophagus cygnaei Mannerheim 1.7 0.3 19 fuscus Erichson 0.2 0.0 2 Curculionidae Carphonotus testaceus Casey 0.3 0.0 3 Cossonus pacificus Van Dyke 0.2 0.0 2 Magdalis alutacea LeConte 0.1 0.0 1 Pissodes fasciatus LeConte 0.2 0.7 7 Pissodes sthatulus (Fabricus) 0 0.1 1 Rhyncolus brunneus Mannerheim 0.2 0.0 2 Rhyncolus macrops Buchanan 1.7 1.6 28 Dermestes talpinus Mannerheim 0.1 0.1 2 Megatoma sp. (cylindnca) (Kirby) 0.6 0.9 12 Megatoma vengatta (Horn) 1.1 0.7 16 Elateridae Ampedus brevis (Van Dyke) 0.7 2.4 24 Ampedus mixtus (miniipennis?) (Herbst) 0 0.3 2

31 Ampedus nigrinus (Herbst) 0.6 0.1 7 Ampedus occidentalis Lane 0.2 0.0 2 Ampedus pullus Germar 0 0.1 1 Athous nigropilis Motschulsky 0.4 0.3 6 Athous rufiventris rufiventris (Eschscholtz) 0.9 2.1 24 Ctenicera aeripennis (Kirby) 0.2 0.0 2 Ctenicera angusticollis (Mannerheim) 0.5 0.9 11 Ctenicera bombycina (Germar) 0 0.1 1 Ctenicera comes (W.J. Brown) 0 0.1 1 Ctenicera hoppingi (Van Dyke) 0.1 0.0 1 Ctenicera lutescens (Fa\\)/sagitticollis (Eschscholtz) 0.2 0.3 4 Ctenicera mendax (LeConte) 0 0.4 3 Ctenicera nebraskensis (Bland) 0.4 1.4 14 Ctenicera nigricollis (Bland) 1.5 1.7 27 Ctenicera pudica (W.J. Brown)+ propola columbiana (Leconte) 5.5 5.1 91 Ctenicera r. resplendens (Eschscholtz) 0.2 0.0 2 Ctenicera semimetallica (Walker) 0 0.1 1 Ctenicera umbricola (Eschscholtz) 0.7 0.4 10 Ctenicera volitans (Eschscholtz) 2.6 0.4 29 Drasterius debilis LeConte 0.5 3.0 26 Eanus sp. #1 0 0.6 4 Negastrius tumescens LeConte 1 0.0 10 Triplax califomica LeConte 0.4 0.7 9 Triplax antica LeConte 0 0.1 1 Triplax dissimulator (Crotch) 0.1 0.1 2 Eucnemidae Epiphanis cornutus Eschscholtz 0.2 0.0 2 Histeridae Paromalus mancus Casey 0.6 0.0 6 Lampyridae Phausis rhombica Fender 0 0.1 1 Lathridiidae Corticaria n. sp. 4.3 2.6 61 Enicmus mendax Fall 0.1 0.3 3 Enicmus tenuicomis LeConte 2.6 2.7 45 Lathridius n. sp. 0.5 0.3 7 Stephostethus breviclavus (Fall) 0.4 0.0 4 Stephostethus liratus (LeConte) 0.9 0.4 12 Agathidium spp. 0.3 0.4 6 Agathidium difformis (LeConte) 0 0.3 2 Agathidium depressum FaW/obtusum Hatch 3.7 2.7 56 Anisotoma globososa Hatch 0 0.4 3 Colon (mylochus) aedeagosum Hatch 0 0.1 1 Colon magnicolle Mannerheim 1.2 0.0 12 Hydnobius sp. # 3 0 0.1 1 Leoides rufipes (Gebler) 0 0.1 1 Triadhron lecontei Horn 0 0.1 1 Lycidae Dyctyopterus spp. 2.3 1.0 30 Emmesa stacesmithi Hatch 0.4 0.0 4

32 Melandrya striata Say 0.1 0.0 1 Phryganophilus collaris LeConte 0.1 0.0 1 Scotochroa basalis LeConte 0.1 0.6 5 Serralopalpus substhatus Haldeman 0.6 0.6 10 Xyleta laevigata (Hellenius) 2.9 0.4 32 Zilora occidentalis Mank 0.2 0.1 3 Melyridae Hoppingiana sp. (hudsonica) (LeConte) 0.2 0.0 2 Trichochrous albertensis Blaisdell 0.4 2.9 24 Mycetophagusdistinctus Hatch 0 0.4 3 Nitidulidae sp # 296 0 0.1 1 Epuraea sp. # 1 0.2 0.3 4 sp.#7 0 0.1 1 sp. # 10 0.1 0.0 1 Epuraea depressa © 1.1 0.0 11 Epuraea flavomaculata Maklin 0.1 0.0 1 Epuraea planulata Erichson 0.1 0.3 3 Eupraea terminalis Mannerheim 0.2 0.1 3 Eupraea truncatella Mannerheim 0.3 0.0 3 Glischrochilus confluentus (Say) 0.1 0.0 1 Omosita discoidea (Fabricius) 3.5 0.3 37 Thalycra mixta H. Howden 0 0.4 3 Oedemerinae Calopus angustus LeConte 1 0.7 15 Pythidae Pytho sp. # 2 0.3 0.1 4 Rhizophagidae Rhizophagus pseudobrunneus Bousquet 0.2 0.0 2 Rhizophagus dimidiatus Mannerheim 2.6 0.1 27 Rhizophagus remotus LeConte 0.6 0.6 10 vindiaeneus Randall 2.1 o.i 22 Scaphidiidae Scaphisoma castaneum Motschulsky 0.1 0.0 1 sp # 328 0 0.1 1 Aphodius fimetarius (Linneaus) 0.8 0.0 8 Aphodius haemorrhoidalis (Linneaus) / pectoralis LeConte 0.1 0.0 1 Aphodius leopardus Horn 0 0.7 5 Cyphonsp.(p) 0.1 0.0 1 Scolytidae Cryphalus ruficollis Hopkins 0.1 0.0 1 Dendroctonus pseudotsugae Hopkins 7699 6.4 77036 Dendroctonus rufipennis (Kirby) 0 0.6 4 Dryocetes affaber (Mannerheim) 0.5 0.6 9 Dryocetes autographus (Ratzeburg) 1.1 0.6 15 Dryocetes betulae Hopkins 0.1 0.0 1 Dryocetes caryi Hopkins/sche/ft' Swaine 0.2 0.0 2 Dryocetes confusus Swaine 0.3 0.1 4 Gnathotrichus retusus LeConte 2.2 0.6 26 Hylastes nigrinus (Mannerheim) 13.2 6.1 175 Hylastes longicollis Swaine 0.3 0.1 4 Hylastes ruber Swaine 5.1 7.3 102 Hylurgops porosus (LeConte) 0 0.1 1

33 Hylurgops rugipennis Mannerheim 0 0.1 1

Ips latidens (LeConte) 0 0.1 1

Ips perturbatus (Eichhoff) 0.1 0.0 1

Ips pini (Say) 0.1 0.0 1

Orthotomicus caelatus (Eichhoff) 0.1 0.0 1 Phloeotnbus lecontei Schedl /picea Swaine 0 0.1 1

Pityogenes hopkinsi Swaine 0.1 0.0 1

Pityogenes plagiatus (LeConte) 0.1 0.0 1 Pityophthorus nitidulus Swainef+ tuberculatus Eschhoff) 0.1 0.3 3

Pityophthorus pseudotsugae Swaine 0.1 0.0 1

Polygraphus convexifrons Wood 0.4 0.1 5

Polygraphus rufipennis (Kirby) 3.2 0.3 34

Pseudohylesinus nebulosus LeConte 4.3 4.7 76 Scierus annectans LeConte 14.9 4.4 180 Scierus pubescens Swaine 0.1 0.0 1

Scolytus sp. (unispinosus) LeConte 0 0.1 1

Scolytus tsugae (Swaine) 1.3 0.0 13

Scolytus unispinosus LeConte 0.4 0.1 5

Trypodendron lineatum (Olivier) 1274 13.0 12832

Trypodendron retusum (LeConte) 2 1.3 29

Trypodendron rufitarsis (Kirby) 0.7 0.3 9 Xylechinus montanus Blackman 5.6 0.7 61 Scraptiidae Anaspis sp. 25.6 1.4 266 Hallomenus sp. 0.1 0.3 3

Orchesia (nr.) castanea 0.1 0.1 2

Sphindidae Odontosphindus clavicornis Casey 0 0.1 1

Staphylinidae Acidota crenata (Fabricius) 0.2 0.3 4

Aleochannae (misc.spp.) 1 0.0 10

Aleochara castaneipennis Mannerheim 0.1 0.0 1

Atheta dentate Bernhauer 0.3 0.1 4 Atrecus macrocephalus (Nordmann) 0.6 0.0 6 Atrecus quadripennis (Casey) 0.1 0.0 1

Bisnius picicornis (Horn) 0.3 0.0 3 Bolitopunctus muncatulus (Hatch) 0.9 3.0 30 Bryophacis Canadensis 0.1 0.0 1

Bryophacis punctulatus (Hatch) 0.1 0.0 1

Carphacis nepigonensis (Bernhauer) 0.4 0.1 5

Dienopteroloma subcostatum (Maklin) 0.1 0.3 3

Earota sp. 1 0.3 12 Eusphalerum spp. (mostly pothos (Mannerheim)) 202 6.9 2068

Hapalaraea sp. #1 0.1 0.0 ' 1

Hapalaraea megarthroides (Fauvel) 0.1 0.4 4

Lathrobium negrum LeConte 0 0.1 1

Leptusa sp. 0.1 0.0 1

Lordithon (Bolitobus) bimaculatus 0.3 0.0 3

34 (Couper) Lordithon cascadensis (Maklin) 0 0.1 1 Lordithon fungicola Campbell 1.2 0.0 12 Megarthrus angulicollis 0.2 0.1 3 Micropeplus laticollis Maklin 0.1 0.1 2 Micetoporus sp. 0.1 0.0 1 Mycetoporus americanus Erichson 0.1 0.1 2 Mycetoporus rufohumoralis Campbell 0.4 1.9 17 Omalium sp. # 1 0.1 0.0 1 Omalium spp. 0 0.1 1 Omalium sp. (foraminosum Maklin) 0.2 0.0 2 Oxytelus fuscipennis Mannerheim 0.2 0.3 4 Pelecomalium testaceum (Mannerheim) 4.5 0.3 47 Philodrepa(?)Dropephylla sp.(nr. Longula Maklin) 0.1 0.4 4 Philonthinii spp. 0.3 0.3 5 Philonthus politus (Linneaus) 0.1 0.0 1 Placusa tacomae Casey 0.1 0.0 1 Pseudopsis sp. 0.1 0.0 1 Quediini spp. 0 0.1 1 Quedius criddlei (Casey) 0.4 0.0 4 Quedius erythrogaster Mannerheim 0,1 0.0 1 Quedius plagiatus Mannerheim 3.5 1.6 46 Quedius rusticus/vilis Smetana 0.3 0.0 3 Quedius s. spelaeus Horn 0.1 0.0 1 Quedius velox Smetana 4.4 4.7 77 Siagonium stacesmithi Hatch 0.6 0.0 6 Staphylinus pleuralis LeConte 0.1 0.1 2 Tachinus basalis Erichson 0.7 0.7 12 Tachinus elongatus Gyllenhal 0.1 0.3 3 Tachinus frigidus Erichson 0.2 0.1 3 Tachinus nigncornis Mannerheim 0.1 0.1 2 Tachinus thruppi Hatch 0.1 0.0 1 Tachypoms sp. 0 0.1 1 Trichophya pilicomis (Gyllenhal) 0 0.1 1 Tenebrionidae Bius estnatus (LeConte) 0 0.3 2 Corticeus praetermissus (Fall) 0.1 0.0 1 Corticeus subopacus (Wallis) 0.2 0.1 3 Eleates explanatus Casey 0.3 0.1 4 Mycetochara fratema (Say) 0.1 0.0 1 Tnbolium audax Halstead 0.3 0.0 3 Abstrulia (nr.) veriegatta Casey 0.9 1.6 20 concolor LeConte 1.9 1.4 29 Throscidae hornii (LeConte) 0.4 0.1 5 Calitys scabra (Thunberg) 0.4 0.4 7 Ostoma ferrugina (Linneaus) 0.6 0.3 8 Thymalus marginicollis Chevrolat 2.7 4.3 57

35 The Venn diagram (Figure 2.3) shows that nearly half (48% or 120/249) of the

analyzed species were common to both baited and control sites: 31% (78

species) were exclusive to baited traps and 20% (51 species) were exclusive to control traps. Table 2.3 shows that 34% of species (86/249) were

represented by a single specimen. Of the 72,546 insects identified, 55,274

(76%) were Douglas-fir beetles (Dendroctonus pseudotsugae).

Figure 2.3. Venn diagram showing the species distribution of flying Coleoptera found in funnel traps baited for Douglas-fir beetle in the Fort St James Forest District, British Columbia. Habitat is Mature/Old growth Douglas-fir (Fd 8, >115 yrs). Data excludes identified necrophagous species or species <1 mm.

Whittaker plots (which display species abundance as average number of

beetles per site by rank) followed logarithmic species distributions for both

baited and control data sets (Figure 2.4). However, pheromone baited sites

contained a greater number of individuals over all (71,477 vs 1,069; baited

and control, respectively), and the pheromone baited samples exhibited a

36 dramatically increased slope for species whose abundance exceeded ten individuals/site (Figure 2.5).

baited control

0.1 -l . Species Rank (1-200)

Figure 2:4. Species abundance of flying Coleoptera expressed by rank as mean abundance per site in preharvest habitat. Abundance is shown for MCOL Seudenol and Frontalin pheromone baited (•) and unbaited/control treatments. Beetles were trapped in Lindgren funnel traps in mature Interior

Douglas-fir (Fd8, >100 yrs) in the Fort St James Forest District, British Columbia.

1 "i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i— «-Nntifl(DNflO(i)OT-N(i)^in

Figure 2.5. Species abundance by rank of the twenty most abundant flying Coleoptera expressed as mean abundance per site in preharvest habitat. Abundance is shown for MCOL Seudenol and Frontalin, pheromone baited (•) and unbaited/control (^treatments. Beetles were trapped in Lindgren

funnel traps in mature Interior Douglas-fir (Fd8, >100 yrs) in the Fort St James Forest District, British Columbia.

37 Species composition of the top 20 most abundant species for baited and

control sites identified 13 species in common (Table 2.4). Rank positions of these species varied between baited and control sites, with the exception of

Eusephalerum pothos: Staphylinidae at rank #3. Differences in mean species

Table 2.4. Twenty most abundant flying beetles species in preharvest baited and unbaited (control) sites. Beetles were trapped in funnel traps baited for Douglas-fir beetle in the Fort St James Forest District, British Columbia.

Preharvest sites = Mature/Old growth Interior Douglas-fir (Fd8, >100 yrs). Listings do not include identified necrophagous species. Twenty Most Abundant Species

Rank #/site Preharvest Baited #/site Preharvest Control t 7890Dendroctonus pseudotsugae Hopkins \7>Trypodendron Lineatum (Olivier) 2 ^QAATrypodendron Lineatum (Olivier) 7Hylastes ruder Swaine (Mannerheim)) 3 285Eusphalerum spp (mostly pothos 7Eusphalerum spp (mostly pothos (Mannerheim)) Mannerheim)) H 163 Thanasimus undatulus (Say) 6Dendroctonus pseudotsugae Hopkins 5 31Anaspis sp GHylastes nigrinus 5Ctenicera pudica (W.J. Brown) +propola .(? 17 Hylastes nigrinus (Mannerheim) columbiana (Leconte) 7 15Scierus annectans LeConte 5Thymalus marginicollis Chevrolat

APolygraphus rufipennis (Kirby) ZAthous rufiventris rufiventris (Eschscholtz) 1 °t AEnicmus tenuicornis LeConte 2Ctenicera nebraskensis (Bland) 20 3 LeConte 2Dyctyopterus spp Megatoma sp (cylindrica) (Kirby)

38 abundance/site between baited and control treatments ranged from one

individual (Quedius velox: Staphylinidae) to 7,877 individuals (Dendroctonus pseudotsugae: Scolytidae).

A total of nine measures of diversity were applied to the data to assess changes in community structure: five measure of species richness (Number of species (S); Margalef (d), Shannon-Wiener (H'-io), Brillouin, and Fisher (

indices), one measure of evenness (Pielou (J') index), one measure of dominance (1-Simpson index (1-D)), and two taxonomic based measures developed by Clarke and Warwick (1999)(Taxonomic diversity (8), and taxonomic distinctness (5 *)).

Results revealed eight out of nine measures were statistically different (at rx =

0.025) between baited and control data. These results include all calculated

richness, evenness, dominance, and taxonomic indices (Figure 2.6, Table

2.5). Species number (S) was the only measure not significantly different

between baited and control data sets, yielding statistically insignificant

averages for baited and control treatments (58.3 and 56.0 species/site

respectively). 1

39 100 s. • • 55

10 - WL * i

1 Value , o • O) 1 - o ! 0.1.-

Calculate d i i c 0 01 s I I I I I

N^ Jyi *2> JL

Diversity Measure/Index

Figure 2.6. Measures of flying beetle diversity from pheromone baited (•) and unbaited/control (a) sites in interior Douglas-fir (Fort St James Forest District, British Columbia). Beetles were trapped in Lindgren funnel traps. Pheromone baited traps were baited with a ternary lure of MCOL, seudenol and frontalin.

Sites were Mature/Old growth Interior Douglas-fir (Fd8, >100 yrs). Indices do not include identified necrophagous species or species < 1 mm.

Despite differences in the types of analysis, their discriminant ability and sensitivity to sample size, all eight calculated diversity indices were statistically different between baited and control treatments. Without exception control sites yielded higher indices of diversity than baited sites

(Table 2.5). In an attempt to isolate the source of the observed differences in species distribution and community diversity, species level non-parametric

statistics were completed through Wilcoxon Rank Sum analysis. Results

identified eight species as having statistically significant differences in mean

abundance (fx = 0.05) between baited and control sites: Six species were

significantly more abundant in baited sites, and two were significantly more

abundant in control sites.

40 Table 2.5. Richness, evenness, and dominance measures of flying beetle diversity for pheromone baited and unbaited preharvest Interior Douglas-fir sites (cx = 0.05). Beetles were trapped in funnel traps baited for Douglas-fir beetle (Mcol, seudenol and frontalin) in the Fort St James Forest District, British

Columbia. Preharvest sites = mature/old growth interior Douglas-fir (Fd8, >100 yrs). Indices do not include identified necrophagous species or species < 1mm. Mean Index Value Mean Index Value T-Test of difference Index per Site - Baited per Site - Control T-Value, P-Value Species Total, S 58.3 + 12.5 56.0 + 8.02 0.30 0.769 (Richness) Margalef, d 6.29 + 1.29 10.9 + 1.35 -4.87 0.000 (Richness) Shannon, H' 0.197 + 0.0861 1.51 + 0.0984 -25.97 0.000 (Richness)

Brillouin 0.444 + 0.198 3.04 + 0.194 -18.37 0.000 (Richness)

Fisher, a 8.39 + 1.94 32.7 + 6.41 -7.12 0.000 (Richness)

Pielou, J' 0.11 + 0.0436 0.87 + 0.0372 -19.75 0.000 (Evenness)

1-Simpson, 1-D 0.194 + 0.126 0.952 + 0.0208 -11.64 0.000 (Dominance) Taxonomic 15.8 + 8.65 67.99 + 1.93 -11.54 0.000 diversity, 8 Taxonomic 61.8 + 6.22 71.5 + 0.772 -3.03 0.023 distinctness, 5*

Species with a greater abundance in baited sites include the pheromone target species - Dendroctonus pseudotsugae (the target Douglas-fir beetle), and other non-target species including: Polygraphus rufipennis (Kirby),

Anaspis sp., Thanasimus undatulus (Say), Rhinosimus viridiaeneus Randall, and Rhizophagus dimidiatus Mannerheim. Species found to have a significantly greater abundance in control sites include Neanthophlax mirificus

(Bland) and Driasterius debilis LeConte. (Table 2.6).

41 Table 2.6. Species of flying Coleoptera statistically more or less abundant between pheromone baited and unbaited control sites in mature to overmature interior Douglas-fir under attack by the Douglas-fir beetle. Pheromone baited traps were baited with a Frontalin, MCOL, Seudenol lure known to aggregate Douglas-fir beetles (Dendroctonus psuedotsugae). Rank Increased Rank Sum abundance Sum Increased abundance Value - Critical Unbaited control Value - Critical Pheromone baited traps Baited Value traps Control Value Dendroctonus pseudotsugae 0 14 Drasterius debilis 11.5 14 SCOLYTIDAE ELATERDIAE Thanasimus undatulus 1 14 eanthophlax mirificus 9.5 • 14 CLERIDAE ERAMBYCIDAE Anaspis sp. 14 14 STAPHYLINIDAE Polygraphus rufipennis 13.5 14 SCOLYTIDAE Rhizophagus dimidiatus 13 14 RHIZOPHAGIDAE Rhinosimus viridiaeneus 13 14 SALPINGIDAE

After identifying species with significant differences in abundance between baited and control sites, these species were removed from diversity data sets, and indices were recalculated. Despite the removal of the eight species, statistically significant differences remained between baited and control data for five of nine measures including two of the four richness measures, as well as all measures of evenness, dominance, and taxonomic diversity.

Subsequent recalculations of Wilcoxon Rank Sum species level analysis found no additional species with significant differences in abundance.

Discussion

Results of this study yield both qualitative and quantitative differences in flying beetle diversity between pheromone baited and unbaited treatments. At the

42 most basic level of analysis, the Venn diagram indicates common and unique species between baited and control data sets. Whittaker plots indicate differences in species distribution between treatments. Non-parametric analysis of abundance at the species level and the parametric analysis of diversity measures at the community level indicate differences in species distribution and abundance between baited and control traps. The data support the rejection of the null hypothesis that pheromone trapping for

Douglas-fir beetles has no impact on the trapping of non-target flying beetle species.

Statistically significant differences observed for 8 out of 9 diversity measures, along with nonparametric analysis of species abundance between baited and control data offer the strongest support for rejecting the null hypothesis.

Diversity, as measured by a calculated index, decreased in all cases with pheromone trapping. Differences observed in indices for richness (d, H',

Brillouin, Fisher's a), evenness (J'), dominance (1/D), Taxonomic diversity (8), and Taxonomic distinctness (5*) indicate decreased species richness, increased dominance, decreased evenness, decreased taxonomic diversity and distinctness in response to pheromone baiting. In an apparent contradiction to the rest of the data, one measure of richness indicated no significant differences between baited and unbaited treatments. Before continuing to assess or discuss the data that rejects the null hypothesis and supports a hypothesis of non-target pheromone influence, it is necessary to

43 address this one analysis that supports the null hypothesis - the analysis of species number (S).

The mean number of species trapped in baited vs unbaited treatments resulted in similar means for species richness (S) per site (Table 2.4). The lack of a significant difference (T-Value = 0.30, P-Value = 0.769 at a = 0.05) in richness is in sharp contrast to all other calculated indices, suggesting that pheromone lures do not change the number of species observed in the data pool. However, closer analysis indicates that this result may be an artefact of low sampling effort. Compared to other indices calculated in the study, S most strongly reflects the species area curve and is highly sensitive to sampling effort (Magurran 1988). The result observed for S is inconsistent with other measures of diversity (including other measures of diversity less sensitive to sampling effort) and does not reflect the change in species composition observed in the Venn diagram (Figure 2.3). However, the result is consistent with total number of species observed for each treatment (Figure

2.3), and the high single species occurrence observed in the study.

The above inferences regarding sampling effort are based on assumptions about the species pool. Defining the total species potential in the context of flying beetles found in Douglas-fir beetle habitat, species manipulation should have occurred from within a pre-existing species pool (of an unknown size) determined by habitat conditions (in this case mature, Douglas-fir beetle

44 attacked, Interior Douglas-fir stands). Despite internal manipulations of distribution and abundance, the total species pool should be finite and similar between baited and control treatments with the exception of transient species.

Based on this assumption, the observed similarity in species number between baited and control replicates is not contradictory with the observed differences in diversity indices. Rather, the current trapping level of 7 sites with 3 traps per site does not appear to encompass the whole of the species area curve.

This result is important because it does not detract from the significant differences in diversity and abundance observed from other calculated indices. It illustrates the need to utilize more than one measure of diversity in data analysis, and limits our ability to draw inferences on community composition and structure to a context of species with greater abundance.

Equal sampling effort maintained for baited and control treatments still allows for comparisons of diversity, distribution and abundance of available species on a relative basis/

Observed differences in measures of diversity suggest that pheromones specific to the Douglas-fir beetle may impact non-target species. But if so, what is the extent of this influence? After removing all eight species with significantly greater abundance in pheromone-baited traps (see Table 2.6) from the data and recalculating diversity indices, significant differences remained in five out of eight calculated indices, indicating that the observed differences in diversity are due in part, but are not strictly owing to, the

45 disproportionate occurrence of those eight species. The results indicate that pheromone trapping for Douglas-fir beetle with this lure impacts a multi- species assemblage of non-target flying beetle species of an undetermined size and species composition. Furthermore, the data suggest the potential presence of kairomonal response at the community level in flying insects associated with the pheromone lure of Douglas-fir beetle. While the presence of a multispecies kairomonal response of this extent has not been published, it is well within the scope of current semiochemical theory.

The Douglas-fir beetle is known to have both positive and negative responses to various host volatiles and conspecific pheromones (Heikkenen and

Hrutfiord 1965). Leading pheromone researchers acknowledge that a large complex of species may join principal scolytids (Borden 1974) acting as secondary exploiters, and decomposers (Huffaker etal. 1984). Odours released from trees during mass attack by bark beetles are thought to influence predators (Borden 1974), competitors, mutualistic species, secondary species, or any combination thereof (Raffa 1991).

One investigation of pheromones and host volatiles associated with southern pine beetles identified fifteen entomophagous, and 13 associate insects

(Dixon and Payne 1980). In the same study, kairomonal action of semiochemicals attracted not only predators, but competitors and less aggressive scolytids. Also in southern pine, chemically mediated behavioural

interactions have been identified in insect colonization sequences, resource

46 partitioning and predation strategies (Birch et al. 1980; Billings and Cameron

1984; Billings 1985, Kohnle and Vite 1984). A recent study on the trapping

efficacy of bark beetle pheromone lures on two Monochamus species

(Cerambycidae) demonstrated a response of the cerambycids to bark beetle

pheromone lures in the absence of host volatiles (Allison et al. 2001). In

interpreting the results investigators suggested that Monochamus spp.

minimised foraging costs by using the pheromones of sympatric beetle

species as kairomones.

By their nature of dispersal, pheromones released into the environment are

available to be interpreted by any other organism that detects them (Birch

1984), and certain chemicals produced by common biosynthetic pathways

may be efficient carriers of specific stereotyped information to more than one

species (Setter and Borden 1992, Huber et al. 2000). The potential for

parsimony of individual semiochemicals across many species means that one

or more chemicals of a pheromone system are available to become part of the

kairomone system of associated species capable of perception and response.

The parsimonious action of frontalin is known to occur for bark beetles

residing in a range of habitats.

Frontalin is known to be associated with at least 12 species of bark beetles in

both aggregation (Table 2.1), and anti-aggregation roles, though in all cases

the pheromone does not produce a maximum species response on its own.

47 Synergistic effects of 2 or more semiochemicals in a volatile mixture disproportionately influences bark beetle behaviour, with the intensity of the effect being dependant on the chemical composition (Libbey et al. 1985), the

nature of reception (Mustaparta 1984), individual semiochemical concentration (Byers 1987), and their geometrical composition (Gries 1992,

Seybold 1993).

When combined with MCOL and seudenol, frontalin triggers an aggregation

response in Dendroctonus pseudostugae (Ross and Daterman 1995). In

Dendroctonus ponderosae, frontalin combined with exo-brevicomin is known to terminate the aggregation response (Pureswaran et al. 2000). In the pheromone systems of Ips grandicollis, frontalin is perceived, but has not been observed to influence the behavioural orientation of the species (Ascoli-

Christensen et al. 1993). Differential responses of bark beetles to parsimonious chemicals appears to result from the ability of dispersing bark beetles to recognize both host and non-host volatiles (Huber et al. 2000) - allowing a beetle species to create a profile of the forest environment during the search for resources that changes context depending on concurrent chemical associations. In the context of this study, the impact of

parsimonious semiochemicals on non-target species warrants consideration,

but the extent of impact of chemical constituents on non-target species is

unknown. Non-target species may be responding to individual components,

binary combinations, or the entire lure.

48 Given that most - if not all - flying beetles will use semiochemical cues (of one type or another, and at one time or another) during their life cycle, it is reasonable to assert that at any one time, within an insect's natural habitat, semiochemicals can have a positive (aggregative), negative (anti- aggregative), or neutral (no apparent effect) effect on an individual or a species. In response to perceived semiochemicals, individual behaviours are altered to varying degrees (or not at all), and the sum of these responses at the individual and species level results in a multispecies, or a community effect.

Proposing a community effect in response to a pheromone lure obliges one to define the nature of such a community. In the context of this study, the community responding to a pheromone lure consists of all species exhibiting a positive response to the Douglas-fir fir beetle pheromone lure, regardless of the extent of the response, or the nature of the association with target species. While it may be more ecologically appropriate to define such a community based on an established relationship with the Douglas-fir beetle, applying any categorical criteria to the data is appropriate only if it can be applied equally to all species. The unknown life histories of majority of species present in the study, and the complexity and range of potential associations between the target and non-target species makes it impossible to determine for all species, the presence or nature of such a relationship

49 (Stephen and Dahlsten 1976). The results indicate a multispecies response to pheromone lures of an unknown extent, and from this result a general concept of community response has been applied to the data - one that contains known, unknown and potential species associations.

Assuming that the Douglas-fir beetle pheromone lure elicits a "community response", then ideally, the community should be identifiable with supporting evidence of species associations in available literature. Of species known respond to Douglas-fir beetle pheromones and lures, literature is available for

2 species: Dendroctonus pseudotsugae, and Thanasimus undatulus.

The most abundant species in pheromone-baited sites was the target species, the Douglas-fir beetle. Not surprisingly this beetle had the greatest increased abundance in baited traps over control traps. The dramatic difference in trap abundance (1,200x increase in baited over control traps) reflects the efficacy of the pheromones previously reported by investigators (Ross and Daterman

1998). The attraction of Thanasimus undatulus (Coleoptera; Cleridae) to

Douglas-fir beetle pheromone components has also been recorded in the literature, so it is not surprising to find this species with significantly greater abundance in baited traps. It is interesting to note, however, that on average the predator/prey ratio of T. undatulus to the Douglas-fir beetle was lower in pheromone-baited traps than in unbaited traps - a condition inconsistent with

50 previous studies evaluating multiple semiochemical lures (Ross and Daterman

1998) or individual semiochemicals (Lindgren 1992).

The remaining four species listed in Table 2.6 with greater abundance in baited or unbaited traps, have no previously published semiochemical response - positive or negative - to Douglas-fir beetle pheromones. In a field of research only 50 years old, the lack of data is not surprising. As of 1990, the pheromone systems of only 100 species of Coleoptera world wide

(primarily Scolytidae) have been investigated (Vite and Baader 1990).

Couple this with a disproportionately low investigation rate in the fields of insect ecology and taxonomy, and it is not surprising that for some species there is no published ecological information at all. While the lack of background makes it difficult to interpret species associations in all cases, data available for a few species allows some insight into the range of potential associations.

In preharvest pheromone trapping, Rhizophagus dimidiatus (Rhizophagidae) was observed to be significantly more abundant in baited traps. The beetle is a known associate of Douglas-fir beetles, occurring as adults and larvae in the galleries of scolytids in (Deyrup and Gara 1978). Its biological association with the Douglas-fir beetle is thought to be as a predator, suggesting a kairomonal response by the rhizophagid occurs to locate

Douglas-fir beetle prey.

51 Rhinosimus viridiaeneus (Salpingidae) is a fungivore associated with the

galleries of the bark beetle Alniphagus aspericollis (LeC). In Alder the beetle

is also found under the bark of broad leaf trees not inhabited by scolytids

(Deyrup and Gara 1978). In this study R. viridiaeneus was significantly more

abundant in baited traps, though a direct association with Douglas-fir beetles

is not presumed. Instead, the feeding preference of this salpingid beetle for

sapwood fungi creates a potential indirect association between the two beetle

species based on minimizing food foraging effort. Assuming the habitat range

off?, viridiaeneus is not limited to broadleaf scolytid galleries/trees, and

includes Douglas-fir as a host, the fungivore could be utilizing Douglas-fir

beetle pheromones as kairomones to locate fresh fungi known to occur in

scolytid galleries.

Polygraphus rufipennis (Scolytidae) is a transcontinental species that occurs throughout western North America. It is a phloeophagus bark beetle restricted

to Abientineae hosts (Bowers et al. 1996), breeding under the bark of smaller

and drier portions of the bole of dead and dieing spruce, lodgepole pine,

limber pine and larch (Furniss and Carolin 1977). Like the preceding

examples, P. rufipennis was also significantly more abundant in pheromone-

baited traps, though again, a direct association with Douglas-fir beetles is not

presumed. The life history of this species is documented to the extent that

52 without further evidence it is unreasonable to suggest a habitat extension that includes Douglas-fir.

Differential trapping may be the result of parsimonious semiochemical influence, though it has not been established whether or not MCOL, seudenol, or frontalin are pheromones of P. rufipennis. The species is known to utilize a terpene based semiochemical (3-methyl-3-butene-1-ol) in conspecific aggregation (Bowers and Borden 1992). The pheromone's functional group structure is similar (albeit different), to that of seudenol, though there's no evidence of a synthesis or interconversional relationship between the two semiochemicals. An alternative interpretation of aggregation can be found in that seudenol is known to part of the aggregation pheromone of the primary attacking spruce beetle (Dendroctonus rufipennis) (Furniss et al. 1976), leading to the potential for P. rufipennis to be 'mistakenly' cross-attracted to a parsimonious pheromone of two, closely related, primary attacking species.

In the absence of parsimony, the species may simply be a tourist - a secondary scolytid common to forest species surrounding Douglas-fir

(namely, spruce and pine) - disproportionately trapped in pheromone baited traps by chance. However, same stand replication (the forest equivalent to match pair design) for 5 out of 7 baited and control sites removes some of the distributional variation that would be required for a significant, random occurrence.

53 In the introduction, 20 non-target species were listed as being associated either with Douglas-fir habitat, one, or more pheromone components of the

Douglas-fir beetle pheromones; or female Douglas-fir beetle frass. Results of this study confirm only one species on that list: Thanasimus undatulus - known to be attracted to frontalin (Lindgren 1992). Of the other ten species known to be attracted to pheromone component(s) or frass, only three species were observed in this study (Dendroctonus rufipennis, Hylastes nigrinus, and

Enoclerus sphegus), and none of these three were found to have abundance levels approaching critical values for either baited or unbaited traps (see table

2.6). Of the remaining eight species known to be attracted to Douglas-fir trees only two species were observed in the study sites. Again, the observed abundances did not significantly differ between baited and control traps, suggesting a habitat association not influenced by the presence of the

Douglas-fir beetle pheromone lure.

Reasons for the observed differences in species responses between previous studies and this one are speculative at best, but worth considering: 1)

Species listed in previous studies, but not observed in this study may have a natural distribution or habitat association that does not extend to the northern limit of Douglas-fir assessed in this study. 2) Species known to be attracted to a single pheromone component may not respond to the combination of chemicals and release rates specific to this study, 3) or conversely, beetles known to respond to the more complex semiochemical bouquet of female

54 beetle frass may not be sufficiently influenced by the relatively simple pheromone lure presented in this study.

The efficacy of pheromone components and their enantiomeric composition appears to be highly fixed in some species (Borden et al. 1980), and variable and labile for others (Lanier and Wood 1975, Herms etal. 1991). In southern British Columbia, Douglas-fir beetles are known to produce an average 45:55 mixture of S-(-)- and R-(+)- MCOL, and will aggregate in response to a racemic synthetic mixture (Lindgren et al. 1992). In the case of frontalin, both coastal and interior Douglas-fir beetle populations will respond to a racemic mixture (though both populations respond to R-(-)- frontalin over the S-(+)-enantiomer) (Lindgren 1992). The effectiveness of racemic

(^MCOL, and frontalin observed in southern and coastal British Columbia is consistent with high number of target species trapped by the 1 frontalin,1

MCOL and seudenol lure of this study at the northern limit of Douglas-fir - suggesting the possibility of a conserved efficacy of pheromone components between spatially separated populations of Douglas-fir beetles. Such a conserved efficacy between populations of Douglas-fir beetles creates the potential for proven pheromone lures to be used as a "marker" against which target and non-target species responses can be measured. However, adapting pheromone traps to assess and monitor multiple species assemblages would require a better understanding of the variation inherent in pheromone trapping, including: a better understanding of the impact of

55 semiochemical composition on regional populations of the target species

(Borden 1994), establishing the ability of pheromone traps to assess beetle

populations, and evaluating the potential long-term impact of pheromone

baiting on both target and non-target species.

Semiochemical composition

As noted in the the methodology, the presence of Seudenol in the pheromone

lure was not a controlled factor. In field conditions, seudenol was observed to

be spontaneously produced from MCOL in the presence of acidic water

(present in release devices from precipitation or condensation of atmospheric water), stabilizing at an equilibrium ratio of 40:60, MCOL:seudenol respectively

(H. Wieser, personal communication1). Because seudenol differs from MCOL

only through the position and configuration of a double bond it will possess an

essentially identical vapour pressure to MCOL, conserving the release rate of the mixure (Carde and Baker 1984). The impact of this interconversion is that

a binary pheromone lure of known enantiomeric composition and release rate

became a ternary pheromone lure, with the third chemical being produced at

an undetermined rate (hours to days), in an undetermined enantiomeric

composition. The presence of this third semiochemical may have resulted in

variable species responses, however, its formation and presence in the lure

did contain elements of predictability and stability.

1 Dr. Helmut Wieser, Department of Chemistry, University of Calgary, AB, Canada.

56 Seudenol is a product of monoterpene oxidation by Dendroctonus bark

beetles (Renwick and Hughes 1975) and has been observed to be produced

by Douglas-fir beetles in a 50:50 and 34:66, S-(-): R-(+) isomeric ratio

(Plummer et al. 1976, and Lindgren et al. 1992 respectively). In combination with MCOL and frontalin, seudenol is synergistic in triggering an aggregation

response in Douglas-fir beetles (Ross and Daterman 1995). Properties of

interconversion between MCOL and seudenol have been observed in the

pheromone production of Dendroctonus frontalis, and are also believed to

occur in D. pseudotsugae as an acid catalyzed rearrangement resulting in a

slightly greater than 50% seudenol, and slightly less than 50% MCOL blend

(Renwick and Hughes 1975). A stable interconversion has also been

observed in spruce beetle field lures (Setter and Borden 1999), and an

approximate 50:50 ratio is consistent with that found in the steam distillate of

D. rufipennis frass (Borden et al. 1996). The formation of seudenol in field

lures was observed to occur within a relatively short time frame following set

up (generally prior to the first collection period of the study (1 week)), and the

interconversion was observed to stabilize at a 60:40, Seudenol: MCOL ratio.

Gas chromatography analysis of field baits collected during and after the

trapping season indicated the presence of both chemicals in tested lures.

These field observations, in addition to the ability of lures to aggregate

Douglas-fir beetle populations throughout the duration of the flight season,

suggests that despite the semiochemical variation (or perhaps because of it)

the lure placed in the Lindgren funnel traps displayed a seasonal efficacy for

57 trapping the target species, against which non-target responses were assessed.

The results of this study were limited to metabolically derived, structurally complex pheromones known to be effective in eliciting a response from the target species. Although a large number of commercially available bark

beetle pheromones (including lures designed for use with Douglas-fir beetles) contain ethanol, this study did not include ethanol in the lure. Ethanol has

been shown to be found in the vascular cambium and transpiration stream of a wide range of deciduous and coniferous trees (MacDonald and Kimmerer

1991), and is attractive to many different species of forest Coleoptera (see

Byers 1992 for a review). This primary aggregant is a synergist with host monoterpenes in attracting a wide variety of forest beetles (including the

Douglas-fir beetle) to baited traps (Pitman et al. 1975, Chenier and Pilogene

1989). The chemical's almost ubiquitous presence in nature, its non-specific source of production (potentially produced by any decomposing organic

material), gives it the ability to generate misleading results in both field and

laboratory studies (Phillips et al. 1988). The potential for non-specific

influence precludes its use in this study. It should be noted that because of the widespread use of ethanol in pheromone efficacy studies, the species

component of ethanol based studies are not directly applicable to the results

of this study (see Peck et al. 1997).

58 Trapping efficacy

In addition to semiochemical variation at the source of dispersal, the impact of

pheromone lures on non-target species is also limited by the ability of traps to disperse pheromones and sample beetle populations. Sampling with unbaited

control traps is thought to result from passive sampling of randomly distributed, and largely common, flying beetles within the forest environment

(Byers et al. 1989), as well as through active sampling in response to the visual profile/colour of the trap (Chenier and Philogene 1989). Pheromone

baited traps sampled not only random and visually mediated sampling, but they appear to include species responses to a variable pheromone plume.

Pheromone plumes from baited traps have been established as a way to

monitor the relative abundance of target scolytids within a defined sampling time and space (Turchin and Odendaal 1996). Aggregation responses from

multi-component pheromones result from beetle responses to the active space of a pheromone plume which is determined by the volatile concentration (emission rate), the extent of overlap between independently

released semiochemical plumes, and species specific behavioural thresholds.

These factors are in turn influenced by changes in ambient temperature, atmospheric conditions (such as sunlight, cloud cover, wind intensity), as well

as geographic features and the presence/extent of forest cover (Elkinton and

Carde 1984). The resulting interaction of pheromones and variable

influences creates an effective sampling area that is elongated, irregular, and

59 of a constantly shifting shape (Turchin and Odendaal 1996). The size of the sampling area is ultimately 3-dimensional, species specific, and only present for those species capable of perceiving and responding to the semiochemical(s) used (Schlyter 1992). Variation in pheromone plume development is present both within a given season, and between seasons. In the study design of this creates the potential for a systematic bias across sampling years (see Table 2.2), however the results suggest that any potential bias was not large enough to overwhelm the impact of the lure.

The extent of variation (resulting from sampling bias or natural factors) in pheromone dispersal and subsequent trapping efficacy is unknown for the species observed in this study, and the relative influence of variables changes depending on scale of measurement. Between sites, environmental variation including geography and forest cover is uncontrolled, but within regional and seasonal variation there likely exists a consistent range of behavioural responses to pheromone lures from a given species. Alternatively, when the context is changed from one species response to one trap catch, it can be assumed that environmental variables and the distribution of the pheromone plume from a single trap or group of traps in a site will vary over time, but be consistent at any given time to the species assemblage within the site. To complicate matters further, interspecific behavioural responses to any given plume may or may not be independent, and may also vary over time and/or environmental factors depending on the physiological condition of the insect

60 (Atkins 1975, Wigglesworth 1984, Salom and McLean 1991). Until further

research is completed at the species level, the inherent variation of natural

systems may seriously influence our ability to accurately assess species

responses over the short and long term.

Long term considerations

The specificity of pheromone blends, and their enantiomeric composition between bark beetles and their predators are considered to be highly co- evolved (Bakke and Kvamme 1981, Payne et al. 1984, Lindgren 1992). In consideration of potential long term effects of pheromone baiting, an alteration of predator-prey interactions resulting from disproportional trapping could theoretically destabilize Douglas-fir beetle populations (Raffa and Klepzig

1989). Spatial heterogeneity inherent in Douglas-fir beetle aggregation patterns, along with behavioural or developmental differences, should theoretically stabilize species interactions (Kareiva 1986, and Raffa 1991 respectively). However, a disproportionate change in predator populations in response to long term synthetic pheromone applications could impact long- term competitive pressure within local beetle populations (Perry 1994), inevitably impacting forest ecosystems at the community/landscape level.

The concern for destabilization goes beyond predator-prey interactions.

Douglas- fir beetles and checkered beetles (Thanasimus undatulus) are predator and prey species impacted by a pheromone lure that, according to

61 our results, impacts an undetermined number of non-target flying beetles. If the responses of the other species are also highly co-evolved-, any species the reproductive success of which is determined by the presence and action of

Douglas-fir beetle pheromone components. If such species secure their reproductive success in response to kairomones present in lures, they may be subject to destabilization from inappropriate pheromone trapping.

Disproportionate trapping of a sex within a species is also a concern. Of non- target species observed to respond to bark beetle attack in southern pine, only females of the genus Xyleborus (Scolytidae) were trapped as the males are reported to be incapable of flight (Dixon and Payne 1980). Sex ratios were not established for the vast majority of species observed in this study, but of the five species with sex ratio data available, two species had exclusively male or female representation: Cossonus pacificus Van Dyke

(Curculionidae) (42 male specimens) and Ischnosoma fimbriatum Cambell

(Staphylinidae) (2 female specimens).

Theoretical concerns of population destabilization warrant consideration, however assessing any potential negative long-term consequences of non- target pheromone trapping also needs to consider the range or extent of pheromone influence on a species, as well as the intensity of baiting relative to the population size, distribution, and available habitat range. Given the current distribution of mature Douglas-fir in British Columbia, pheromone

62 sampling may have no measurable long term impact on species diversity at this time. However, increasing losses of old growth stands resulting from current harvest levels, and an increase in the intensity of bark beetle management efforts in recent years warrants some consideration of potential long term impacts of mass trapping with synthetic lures.

Summary

Based on the results of this study, pheromone lures known to create an aggregation response in Douglas-fir differentially influence a multi-species assemblage of non-target flying beetles. The study identified six previously

unassociated species potentially aggregated - as well as two species potentially repelled (antiaggregated) by Douglas-fir beetle pheromones (Table

2.6). Results also indicate that the non-target pheromone influence is not

limited to significantly abundant species. However, the data provide us no clue about the extent of lure influence, or the exact nature of species

associations. Ideally, a combination of life history knowledge combined with

behavioural and physiological studies could provide insight into the reason(s) for both pheromone and habitat association, but such information is

unavailable for the vast majority of species.

Pheromones are complex. Their complexity, and potential for highly specific

use make them one of the most promising tools available for the management

of economically damaging insects. However, the inherent integration of

63 pheromones and natural systems, means that their use as a management tool

may not be species specific, and may instead result in beetle management at

a community level. The impact of Douglas-fir beetle pheromone lures on non-

target species identified in this study indicates that in this circumstance - there is a need for further investigation on the impact of pheromone lures as a

management tool.

64 CHAPTER III Impact of Harvesting on Pheromone Biased Diversity Sampling

Introduction

The previous chapter reported that funnel traps baited with Douglas-fir beetle pheromones MCOL, seudenol, and frontalin result in trap catches of flying beetles different from those caught in unbaited traps in beetle attacked, mature to overmature Douglas-fir habitat. Changes observed in diversity

indices and species abundance patterns describe a phenomenon of non- target response to synthetic pheromone lures. However, these results give no indication of the extent of the semiochemical attraction, nor do they provide any insight into the effect of Douglas-fir beetle pheromones in other habitats successfully utilized by the Douglas-fir beetle.

Pheromone systems of Dendroctonus beetles allow epidemic and endemic

populations to successfully exploit their host in response to a wide range of

internally (endogenously) and externally (exogenously) initiated disturbance events (Wood 1982; Shore etal. 1999 ). Under non-catastrophic disturbance conditions, attack from Douglas-fir beetles is closely associated with patch mortality and gap formation in mature stands (Lewis and Lindgren

2000). Forest harvesting is one type of small-scale disturbance event known to aggregate Douglas-fir beetles (Lejeune et al. 1961). In an ironic twist for forest managers, the growth of bark beetle populations in the interior of British

Columbia can often be traced back to logging disturbance designed to remove

65 active beetle populations - in part because successful breeding occurs in

logging residue (slash, stumps, or coarse woody debris) in the seasons

immediately following harvesting (Lejeune et al. 1961). Postharvest

pheromone baiting is one method used to manipulate, monitor, and reduce

local Douglas-fir beetle populations (Ross and Daterman 1997, Guthrie and

Wieser 1997). However, the combined impact of harvesting and pheromone

baiting on non-target flying beetles is unknown.

Harvesting trees infested with bark beetles, as a method of beetle population

control, changes forest structure and composition (Paulson 1995).

Harvesting practices of clear-cut, or patch-cut logging in North American temperate forest are known to effect change in the diversity and abundance of

plants, mammals, birds, amphibians, and insects (Perry 1994, Karr and

Freemark 1985, Seip 1996, Bury and Corn 1988, and Schowalter 1985

respectively). It is generally accepted that a trend of increasing diversity

occurs with secondary forest succession (Perry 1994). The exact

composition, structure, and rate of diversity increase is dependent on

management intensity and the local species pool (Oliver and Larson 1990,

Perry 1994, Lundquist 1995). Within insect assemblages, long term trends

are thought to reflect functional responses to changing vegetation (Schowalter

1985), while short term changes associated with successional events are

usually defined in the context of insect associations with coarse woody debris

(CWD) (Heliovaara and Vaisanen 1984).

66 A large number of insects are known to utilize dead or dying trees (both standing and fallen) for food, protection, accomodation, reproduction, or combinations thereof (Harmon et al. 1986). Bark beetles (Scolytidae), and wood borers (Cerambycidae and Buprestidae) are the most commonly described families associated with stage one (0-6 years) of tree death and decay (Harmon et al. 1986, Knight and Heikkenen 1980, Caza 1993, and

Dajoz 2000), though the above general descriptions are thought to seriously under-represent the species (and family groups) observed in association with stage one decay of CWD and beetle attacked standing (dying) trees. At least

30 species of Coleoptera from 12 families have been observed to be associated with recently-felled spruce (Gara etal. 1995), 61 species from 25 families have been associated with beetle-attacked, dying pine (Stephen and

Dahlsten 1976), and 86 species from 26 families have been associated with mountain pine beetle-attacked lodgepole pine, yellow pine and western white pine (De leon 1934).

Studies assessing long term insect changes associated with harvesting consist of a relatively small number of studies - most of which document changes in populations (Coleoptera: Carabidae). Overall carabid biodiversity tends to increase with regeneration associated with

secondary succession of mature temperate forests (Lenski 1982, Halme and

Niemela 1993, Vaisanen et al. 1993, Niemela et al. 1992, McDowell 1998,

67 Lavallee 1999). Individual species responses to harvesting practices are thought to depend on the ability of the species to adapt to the successional

habitat (Werner and Raffa 2000).

The Douglas-fir beetle is a primary attacking bark beetle that utilizes a dynamic pheromone system to locate, colonise, and utilize dead and dying

Douglas-fir (Pitman and Vite 1974). The synthetic reproduction and simultaneous release of the aggregation pheromones MCOI, seudenol, and frontalin in a lure is known to trigger an aggregation response in Douglas-fir

beetle populations (Ross and Daterman 1998), and can be used to trap

and/or monitor beetle populations in preharvest and postharvest conditions

(Borden 1994). Prior to the Fort St James Douglas-fir beetle research

project, and this resulting thesis, the combined effect of harvesting and

pheromone trapping on non-target flying beetles had not been reported in the

literature.

Pheromone baiting was assessed under preharvest conditions and up to five years following harvesting for beetle attack (a duration of trapping designed to

encompass the aggregation period of Douglas-fir beetles following harvesting, within the 0-6 year, first stage of Douglas-fir decay). The null hypothesis was that after harvesting, pheromone baiting would have no impact on the flying

beetle community beyond the target species.

68 Methods

Flying beetles were gathered from 65 seasonal data sets recorded from 21

sites in the Fort St James Forest District, Fort St James, British Columbia,

between 1994 and 1997 (Table 3.1). A minimum of 11 sets of seasonal data were gathered from each of old-growth (10 baited, 7 control replicates), first

season (11, 1), second (11, 1), third (10, 5) and fourth/fifth season (6,5)

postharvest condition (baited, control replication respectively for all

treatments).

Table 3.1. Site list with harvest stage, trapping year, and biogeoclimatic classification for pheromone-baited and unbaited sites in the Fort St James Forest District, British Columbia. Subzone definitions can be found in Meidinger and Pojoar (1991). Biogeo• Preharvest Post-1 Post-2 Post-3 Post -4/5 Site Name climatic Bait Control Bait Control Bait Control Bait Control Bait Control Subzone 1 TachieHill SBSdw3 94 95 96 97 97 2 Tachi-Pinchi SBSdw3 94 97 95 96 97 3 GP 8 & 9 km SBSmk 94 97 97 4 GP 13 km S SBSmk 94 95 96 97 97 5 GP 13 km N SBSmk 96 97 6 APC/CP18 SBSdk 97 96 97 7 Hobson Is. SBSdw3 94 95 96 97 97 8 AP1-4 SBSdw3 95 96 97 97 96 97 97 9 WD SBSdw 97 97 95 97 97 10 Siesmic SBSdk 97 11 RNE SBSk3 97 97 12 RNW SBSk3 96 97 13 Pinchi Hill SBSd06 97 97 14 CP 123 96 97 97 15 Kuz Che SBSwk3 96 16 CP 32-80 96 17 CP 46-222 SBSwk3 96 97 97 18 CP 115-1 SBSwk3 96 96 97 97 19 115 (Randy) SBSwk3 97 97 20 CP 118 SBSwk3 96 97 97 21 CP120 SBSdw3 96, 97 97

69 Sites were monitored from one year to a maximum of five consecutive years.

At all sites, 12-funnel Lindgren funnel traps (Phero Tech Inc., Delta, B.C.) were used for both pheromone-baited and unbaited (control) flight

interception. Trap protocol required that a standard minimum of four traps per

site be placed at least 50m apart and at least 50m inside the habitat margin.

Traps were placed so that collection cups were suspended 1 -1.5 m above

ground, clear from interference from vegetation (see Figure 2.2, Chapter 2).

All collection cups contained a 3cm2 piece of neuro-insecticide impregnated

plastic to prevent insect escape, and reduce necrophage activity. Collection

cups released rainwater through a bottom screen to create a dry trapping

system. No British Columbia Resource Inventory Committee (RIC) standards

are available for this sampling technique.

Traps were placed within preharvest or recently harvested stands with >80%

Douglas-fir ( Fd8) composition, in four Biogeoclimatic subzones: SBSdw,

SBSdk, SBSwk, & SBSmk (Meidinger and Pojoar 1991) (seeTable 3.1).

Stand age prior to harvesting was mature to over mature (110 - 350 years).

The trapping period covered the flight season of the Douglas-fir beetle. Sites

were initiated in late April/early May, and were maintained until mid to late

August. The timing of trap set-up was determined by climate and site factors

including snow pack and road conditions. All sites were implemented prior to

70 the onset of the Douglas-fir beetle flight season. Traps were removed after

both flight peaks of the Douglas-fir beetle had passed and field personnel

observed two-three weeks of low to no Douglas-fir beetle numbers at all sites.

Trap samples were collected weekly, bimonthly, or monthly as determined by

schedule or site accessibility.

Pheromone traps were baited with a Douglas-fir beetle aggregation lure

developed by researchers at the University of Calgary. The lure consists of a

ternary blend of racemic (*) frontalin (Fn) (1,5,-dimethyl-6,8-

dioxabicyclo[3.2.1]octane), racemic i1) MCOL (1-methylcyclohex-2-enol) and

seudenol (3-methylcyclohex-2-en-1-ol) of an undetermined enantiomeric

composition. The release rate of frontalin was independently regulated from

the release rate of MCOL and seudenol. The release rate of1 Fn was 0.3

mg/day from capillary tubes of 1.0 mm diameter. The MCOL-seudenol blend was achieved by dispensing pure1 MCOL at an average rate of 3.0 mg/day

from a microcenterfuge tube with a 2mm opening in the cap. The open

system of dispersal for MCOL allowed atmospheric water (-OH) (thought to

result from condensation) into release devices, creating an aqueous solution

and an MCOL-seudenol interconversion of an undetermined rate (hours-days)

that was observed to stabilize at a 40-60 ratio (respectively). The resulting

ternary lure was placed in a hooded cradle attached inside the third lowest

funnel of the Lindgren trap (Figure 2.2, Chapter 2). Lures contained enough

semiochemical for the duration of the trapping season and were only changed

71 in response to animal damage, or due to random selection for gas chromatography analysis (to monitor chemical integrity).

Following collection, samples were bagged and frozen to minimize desiccation. Samples remained in frozen storage until shipment to the

University of Calgary for the first of two sorting procedures. From frozen storage, samples were soaked in 70% ethanol for a minimum of 30 minutes and strained using a 1mm wire mesh. Samples were then dried at room temperature in a fume hood from one to four hours (time dependent on sample size), and hand sorted with the aid of dissecting scopes. This initial sort separated out the target species (the Douglas-fir beetle) from other

Coleoptera and removed obvious debris from the samples. Target species abundance was estimated by weight through regression analysis (Appendix II) and target species identification was achieved through either census or sampling depending on the number of beetles contained within the sample.

For samples with 100 or less Dendroctonus beetles, identifications were achieved through census, while identification of samples with greater than 100

Dendroctonus beetles was achieved through a sub sampling protocol.

Samples containing greater than 100 Dendroctonus specimens were themselves sampled for species composition by identifying the first 100

Dendroctonus beetles sorted from the sample. Non-target species were re- frozen and transported to the University of British Columbia for final sorting, mounting, and identification.

72 Beetle identifications were accomplished by the use of available keys, and by

reference to named specimens in both the Spencer Entomological Museum

(University of British Columbia) and the Canadian National Collection of

Insects (Agriculture and Agri-Food Canada, Ottawa). Species groups were

identified through the assistance of taxonomic specialists in Canada, the

United States, and New Zealand. Appendix I lists specialists and their

assistance to this project.

Following identification of all specimens, collection and sample data were entered into an MSExcel spreadsheet. Prior to analysis, data were reduced to

a standard of three traps per site. Trap selection was determined by missing

sample (eg. broken/damaged trap) or by random selection. Data editing was

limited to the removal of necrophagus species suspected to be a direct artifact

of the trapping process (reference), or a species the abundance of which may

be altered as the result of processing protocols (such as the inclusion of

species smaller than the 1mm mesh size used in sample cleaning). Editing was limited to this criterion because the objective of the study was to assess

all non-target species potentially influence by pheromone trapping.

Data analysis included four areas of assessment outlined here and fully

described below: 1) Whittaker rank distribution of species abundance within

treatments (successional years identified as preharvest, 1st season

73 postharvest, 2 , 3 , & 4/5 season postharvest). 2) Calculation of diversity indices (richness, evenness, dominance and taxonomic) within treatments for comparison between treatments. 3) Species abundance comparisons

(Wilcoxon Rank-Sum analysis) between baited and control data within each treatment, and 4) an assessment of trends in abundance for individual species between treatments.

Species distribution

Species distribution analysis within treatments consisted of Whittaker (rank abundance) plots for all available data from all treatments: 10,7 preharvest

(baited, control replicates respectively); 11,1 First season postharvest (post-

1); 11,1 post-2; 10,5 post-3; and 5,5 post-4/5 (Table 3.2). To allow for between treatment comparisons, all Whittaker plots were calculated by the average abundance for each species/site.

Table 3.2. Site replication of baited and control data across treatment years for the Fort St James pheromone study. * Indicates sites in which diversity calculations are included for general comparison only. Treatment Preharvest Post 1 Post 2 Post 3 Post 4/5 year Baited or bait control bait control bait control bait control bait control control # of site replicates 10 7 11 1* 11 1* 10 5 6 5

74 Diversity analysis

A total of 9 measures of diversity were applied to the data to assess richness, evenness, dominance and taxonomic diversity. Analysis was run on a five-

replicate subset of preharvest data conisistent with available replication for postharvest conditions. The resulting low, but equal, sampling effort allowed the greatest amount of cross treatment comparisons, with the largest possible data set, without compromising the requirements of equal sample size

inherent in biodiversity analysis.

Low sampling effort with replication was achieved by limiting the sampling unit to a single site composed of three traps. The five site replicates of baited and control traps for each of the five treatment years, allowed for statistical comparisons of mean index values. Analysis consisted of 42 sites (Table

3.3): five sites for each of preharvest, 1st, 2nd, 3rd, and 4th/5th season

postharvest conditions and five control sites for each of preharvest, 3rd, and

4th/5th season postharvest conditions. Single control replicates for 1st, and 2nd season postharvest sites were calculated and included in data analysis for general comparison only.

A total of nine measures of diversity were applied to all data to assess

changes in community structure: five measures of species richness (Number

of species (S); Margalef (d), Shannon-Wiener (H'i0), Brillouin, and Fisher (a)

indices), one measure of evenness (Pielou (J1) index), one measure of

75 dominance (Simpson index (1- )), and two recently developed taxonomic based measures developed by Clarke and Warwick (1998) (Taxonomic diversity (8), and taxonomic distinctness (8*)). Richness and biodiversity indices were calculated using the software program - PRIMER. Within treatment comparisons of baited and control indices were subject to statistical analysis by a T-test for difference = 0 (vs not = 0) using the software program

MINITAB 2002.

Table 3.3. Site list for 5-replicate diversity analysis. Site data includes harvest stage, trapping year, and biogeoclimatic classification (Fort St James Forest District, B. C). Site Name Biogeo• Preharvest Post -1 Post -2 Post -3 Post -4/5 climatic Bait Control Bait Control Bait Control Bait Control Bait Control Subzone Tachie HII SBSdw3 94 95 97 97 Tachi-Pinchi SBSdw3 94 97 95 96 GP 8 & 9 km SBSmk 94 97 97 GP 13kmS SBSmk 94 95 96 97 97 Hobson Is. SBSdw3 94 95 97 97 AP1-4 SBSdw3 97 97 97 97 WD SBSdw3 97 97 97 97 RNE SBSk3 97 97 Pinchi Hill SBSd06 97 97 CP 123 96 97 97 CP 46-222 SBSwk3 97 CP 115-1 SBSwk3 96 97 CP118 SBSwk3 96 97 CP120 SBSdw3 96 97

Species abundance - baited vs. control

Abundance assessment at the species level consisted of a baited and control

comparison of mean abundance for all species, in all sites (with all available

data) for Preharvest, Post 3 and Post 4/5 conditions. Species data were

76 subject to non-parametric statistical analysis by Wilcoxon Rank-sum test.

Harvest years with only one control replicate (1st and 2nd season postharvest) were omitted from this analysis.

Species trends

Using baited and unbaited data sets, mean species abundance was recorded from preharvest, and 1st, 2nd, 3rd and 4/5th season postharvest conditions.

Based on the observed change in mean abundance/site across treatments, species were then classified by trend: Increasing abundance, decreasing

abundance, increasing then decreasing, decreasing then increasing, or no

observed trend. Species with single occurrences were omitted from analysis.

Category determination was based on a trend being present for at least four out of five data points, with the outlying data point (when present) occurring

not more than one rank position outside of expectation. Individual species abundances across treatment years were then added to produce single trend

lines (where abundance is presented as the sum of mean abundances/site for species within the trend).

Results

A total of 490,028 individuals from 512 identified species and 129

recognizable species groups (RTU's)/morphospecies from 67 families were

identified in preharvest and postharvest baited and control sites. Out of this

77 species complement, four species (Nicrophorus defodiens Mannerheim,

Nicrophorus guttula Motschulsky, Nicrophorus investigator Zetterstedt,

(SILPHIDAE); and Catops egenus (LEOIDIDAE)) representing 5,436

individuals were identified as necrophagus and were removed from data. An

additional 13 species (C/'s striolatus Casey, Cryptophorus sp., Dolichecis

indistinctus Hatch (CIIDAE); Atomaria sp. 1, (3,4&5),8, (CRYPTOPHAGIDAE);

Corticaria gibbosa (Herbst), and Lathridius hirtus Gyllenhal, Melanopthalma

americana (LATHRIDIIDAE); Crypturgus borealis Swaine (SCOLYTIDAE);

Micropeplus tesserula Curtis, Phloenomus lapponicus (Zetterstedt), Stenus

assequens Rey (STAPHYLINIDAE)) representing 364 individuals with body

size <1mm were removed because of sampling inefficiencies. This left

484,228 individuals from 624 species/RTU's for data analysis (Table 3.4, 3.5,

Appendix III).

78 Table 3.4. Summary results of abundance and species number of flying beetles occurring in baited and unbaited Lindgren funnel traps in mature and recently harvested Douglas-fir habitat (1,2,3,4/5 years postharvest) (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle {Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin. Treatment # Site Total # of Total # of Total # # of non- replicates species beetles Douglas-fir target (3 trap/site) beetles species Preharvest 10 Baited 209 94,921 76,991 17,930

7 Control 171 1,069 45 1,024

Post 1 11 Baited 416 198,385 141,452 56,933

1 Control 152 2,285 16 2,269

Post 2 11 Baited 413 128,728 113,984 14,744

1 Control 113 480 15 465

Post 3 10 Baited 350 45,983 41,528 4,455

5 Control 228 1,431 38 1,393

Post 4/5 6 Baited 262 9,258 6,824 2,434

5 Control 231 1,779 10 1,769

All sites 65 625 484,319 380,903 103,415

79 Table 3.5. Summary results of mean abundance per site and total abundance of flyng beetles occurring in baited and unbaited Lindgren funnel traps in Mature and recently harvested Douglas-fir habitat (1,2,3,4/5 years postharvest) (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, Seudenol, & Frontalin. Species with a mean abundance/site >100 have been rounded to the nearest whole number.

B •£ B B B B (/) 03 -5? 'co ro t c "c "c CD ^- roCD E. roCD roCD roCD E p CD CD CD E, E, E, E E E o T3 CD TJ T3 "O C Q. B B ro O CL in 'ro 'ro 'ro o o o CD .O -0 to o ID ro n o ro2 CM m CM ?f .c to to to CO to CD 3 CD o o o o o O o O o Family Genus species CL CL CL CL 0- CL CL CL CL O. r- Alleculidae Isomera (nr) comstoki Papp 0.0 0.1 0.00 0.0 0.0 0.0 0.0 0.0 000 1

Anobiidae sp. # 1 0.0 0.0 0.00 0.0 0.0 0.0 0.0 0.0 0.0.1 1 Caenocara scymnoides LeConte 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Desmatogaster subconnata (Fall) 0.0 0.0 0.0 0.1 0.2 0.0 0.0 1.0 0.0 0.0 Dorcatoma (prob) americana # 87 0.4 0.0 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 6 Ernobius gentilis Fall 0.0 0.4 0.0 0.0 0.2 0.0 1.0 0.0 0.0 0.6 9 Ernobius nigrans Fall 0.0 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 3 Hadrobregmus americanus (Fall) 0.0 0.3 0.5 1.1 0.0 0.0 0.0 0.0 0.2 1.4 28 Hadrobregmus quadrulus (LeConte) 0.0 0.0 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Hemicoelus carinatus (Say) 0.0 0.1 0.1 0.2 0.2 0.1 0.0 0.0 0.2 0.0 7 Microbregma e. emarginatum (Duftschmid) 0.6 2.3 2.4 0.7 0.0 0.7 1.0 6.0 0.4 0.2 79 Ptilinus lobatus Casey /basalis Leconte 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1

Utobium elegans (Horn) 0.1 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Xyletinus rotundicollis R.E. White 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 4 Allandrus populi Pierce 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Tropideres fasciatus (Olivier) 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 Bostrichidae Stephanopachys substriatus (Paykull) 0.0 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Buprestidae Anthaxia inornata (Randall) 0.2 1.7 0.5 0.3 0.0 0.1 0.0 1.0 0.2 0.0 32

Buperstis langi Mannerheim 0.0 0.0 0.0 0.2 0.3 0.0 0.0 0.0 0.0 0.4 6

Buprestis lyrata Casey 0.0 6.5 4.5 4.5 6.2 0.0 0.0 11.0 8.4 7.2 291 Buprestis nuttalli Kirby 0.0 0.2 0.7 0.3 1.0 0.0 0.0 0.0 0.0 0.6 22 Chrysobothris carinipennis LeConte 0.0 0.0 0.3 0.4 0.0 0.0 0.0 0.0 0.6 0.0 10

Dicerca tenebrica (Kirby) 0.0 2.2 9.4 9.6 7.5 0.0 1.0 3.0 12.0 8.8 376 Dicerca tenebrosa (Kirby) 0.0 3.3 3.0 0.9 1.0 0.0 5.0 0.0 1.6 0.4 99 Melanophila drummondi (Kirby) 0.0 0.5 0.4 0.2 0.0 0.0 1.0 0.0 0.4 0.0 14

Byrrhidae Curimopsis sp. 0.1 0.0 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.2 4

Bhyrrhus sp. 0.0 0.1 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 3 Cytilus sp. (alternatus) (Say) 0.0 0.3 0.5 0.2 0.7 0.0 1.0 0.0 0.0 0.2 17

Byturidae Byturus unicolor Say 0.0 0.0 0.2 0.1 0.3 0.0 1.0 0.0 0.2 0.0 7

80 Cantharidae Malthodes sp. 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.2 0.0 2 Podabrus fissilis Fall 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Podabrus piniphilus (Eschscholtz) 0.1 0.4 0.5 0.1 0.3 0.6 3.0 0.0 0.4 0.6 25 Podabrus scaber LeConte 0.0 0.0 0.2 0.0 0.2 0.0 2.0 0.0 0.0 0.2 6 Podabrus sp. # 613 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Silis d. difficilis LeConte 0.1 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.2 0.0 4 Carabidae Agonum placidum (Say) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Amara apricaria (Paykull) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 Amara discors Kirby 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Amara erratica (Dutschmid) 0.0 1.2 0.5 1.0 0.2 0.0 1.0 1.0 0.6 0.6 37 Amara idahoana (Casey) 0.1 0.0 0.1 0.1 0.2 0.0 0.0 0.0 0.0 0.0 4 Amara familiaris (Dutschmid) 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Amara latior (Kirby) 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Amara laevipennis Kirby 0.0 0.4 0.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 11 Amara Httoralis Mannerheim 0.0 0.3 0.5 1.2 0.3 0.0 0.0 0.0 2.2 0.0 33 Amara lunicollis Schodte 0.0 0.5 0.6 0.8 0.5 0.0 0.0 0.0 0.0 0.4 25 Amara patruelis Dejean 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Amara sinuosa (Casey) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Bembidion canadianum Casey 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Bembidion forestriatum (Mutschulsky) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Bembidion grapii Gyllenhal 0.0 0.8 0.6 0.4 0.0 0.0 0.0 0.0 0.2 0.4 23 Bembidion nigripes (Kirby) 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Bembidion quadrimaculatum (LeConte) 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Bembidion tetracolum Say 0.0 0.1 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 5 Bembidion timidum (LeConte) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Bembidion versicolor (LeConte) 0.0 0.0 0.3 0.2 0.0 0.0 0.0 0.0 0.0 0.0 5 Bradycellus congener (LeConte) 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.2 3 Bradycellus lecontei Csiki 0.0 0.3 0.2 0.8 0.3 0.0 0.0 0.0 0.0 0.4 17 Bradycellus neglectus (LeConte) 0.0 0.4 0.5 0.5 0.7 0.0 0.0 0.0 0.0 0.2 20 Bradycellus nigrinus (Dejean) 0.1 0.5 1.1 2.5 1.8 0.0 0.0 0.0 1.2 5.8 90 Calathus advena (Leconte) 0.1 0.0 0.3 0.0 0.2 0.1 0.0 0.0 0.0 0.0 6 americanus Dejean 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Elaphrus clairvillei Kirby 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 2 Harpalus animosus Casey 0.0 0.0 0.0 1.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Harpalus fuscipalpis Sturm 0.0 0.0 0.2 0.0 0.2 0.0 0.0 0.0 0.0 0.0 3 Harpalus laevipes Zetterstedt 0.0 0.1 0.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6 Harpalus nigritarsis CR. Sahlberg 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Harpalus opacipennis (Haldeman) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Harpalus obnixus Casey 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Harpalus somnulentus Dejean 0.0 0.1 0.5 0.2 0.0 0.0 0.0 0.0 0.6 0.2 13 moesta LeConte 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Notiophilus aquaticus (Linneaus) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Notiophilus directus Casey 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1

81 Pterostichus adstrictus Eschscholtz 0.0 0.1 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 3 Sericoda quadripunctata (DeGeer) 0.1 1.2 0.5 0.6 0.0 0.0 0.0 0.0 0.0 0.2 27 Stenolophus fuliginosus Dejean 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Syntomus americanus (Dejean) 0.1 0.6 0.5 0.5 0.3 0.0 0.0 0.0 0.2 0.2 22 Synuchus impunctatus (Say) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Tachyta angulata Casey 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Tachyta rana (Casey/Say) 0.0 0.0 0.0 0.1 0.2 0.0 0.0 0.0 0.0 0.0 2 Trachypachus holmbergi Mannerheim 0.0 2.2 1.3 0.1 0.2 0.0 1.0 0.0 0.2 0.0 42 Trichocellus cognatus (Gyllenhal) 0.0 1.8 1.9 2.1 1.3 0.0 0.0 2.0 0.6 0.6 78 Cephaloidae Cephaloon tenuicorne LeConte 0.3 1.2 1.9 1.1 0.7 0.6 0.0 2.0 0.2 1.0 64 Cerambycidae Acmaeops p. proteus (Kirby) 0.0 0.4 0.5 0.2 0.3 0.0 1.0 0.0 0.0 0.2 16 Asemum striatum (Linneaus) 0.1 5.1 0.3 0.0 0.0 0.1 8.0 0.0 0.0 0.0 69 Callidium cicatricosum Mannerheim 0.0 0.1 0.0 0.1 0.2 0.0 0.0 0.0 0.2 0.2 5 Clytus sp. 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Cortodera sp. # 514 0.0 0.0 0.2 0.2 0.2 0.0 0.0 5.0 0.4 0.2 13 Corcodera (prob) longicornis (Kirby) 0.0 0.2 0.1 0.3 0.2 0.1 0.0 0.0 0.0 0.2 9 Cortodera m. militaris (LeConte) 0.4 1.5 1.5 0.3 0.2 0.0 4.0 0.0 0.0 0.6 48 Cosmosalia chrysocoma (Kirby) 0.0 0.1 0.1 0.2 0.2 0.0 0.0 0.0 0.4 0.0 7 Dicentrus bluthneri LeConte 0.2 0.2 0.1 0.0 0.0 0.3 0.0 0.0 0.0 0.0 7 Evodinus monticola vancouveri Casey 2.7 0.2 0.0 0.1 0.2 0.9 3.0 1.0 0.0 0.0 41 Gnathacmaeops pratensis (Laicharting) 0.0 0.3 0.4 0.1 0.0 0.0 0.0 0.0 0.0 0.0 8 Grammoptera subargentata (Kirby) 0.0 0.4 0.1 0.7 0.2 0.1 0.0 1.0 0.4 1.2 23 Judolia m. montivagens (Couper) 0.0 0.4 2.5 0.7 0.3 0.1 1.0 10.0 0.4 0.6 58 Megasemum asperum (LeConte) 0.1 0.3 0.8 0.6 0.8 0.6 3.0 1.0 0.4 0.8 41 Monochamus spp. 0.0 1.4 0.9 0.3 0.3 0.0 4.0 0.0 0.2 0.0 35 Neanthophlax mirificus (Bland) 0.3 2.8 1.4 1.4 5.7 1.7 17.0 3.0 4.6 5.0 177 Neoclytus m. muricatulus (Kirby) 0.0 0.1 0.1 0.4 0.3 0.0 0.0 0.0 0.6 0.2 12 Pachyta lamed liturata Kirby 0.0 1.9 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 25 Pidonia scripta (LeConte) 0.5 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 6 Phymatodes dimidiatus (Kirby) 0.5 0.6 0.5 0.2 0.5 0.1 0.0 1.0 0.0 0.0 24 Phymatodes (nr.) fulgidus Hopping 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.0 0.0 0.0 1 Phymatodes maculicollis LeConte 0.0 0.0 0.0 0.0 0.0 0.1 0.0 q.o 0.0 0.0 1 Pogonocherus mixius Haldeman 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Pogonocherus penicillatus LeConte 0.0 1.5 0.3 0.0 0.0 0.0 11.0 0.0 0.0 0.0 30 Poliaenus oregonus (LeConte) 0.0 0.2 0.1 0.0 0.2 0.0 1.0 1.0 0.0 0.0 6 Pygoleptura n. nigrella (Say) 0.1 0.6 0.5 0.9 0.7 0.0 0.0 0.0 0.6 0.8 34

82 Rhagium inquisitor (Linneaus) 0.0 9.7 10.4 1.9 0.3 0.0 9.0 11.0 0.4 0.2 265 Semanotus ligneus (Casey) 0.0 0.1 0.0 0.0 0.0 0.0 1.0 0.0 0.0 0.2 3 Spondylis upiformis Mannerheim 4.3 3.1 0.7 1.0 1.7 3.3 4.0 0.0 1.0 0.6 140 Strictoleptura canadensis cribripennis (LeConte) 0.1 3.6 6.6 13.1 8.3 0.0 0.0 1.0 12.6 5.6 388 Tetropium velutinum LeConte 0.4 1.5 0.6 0.1 0.0 1.3 2.0 1.0 0.0 0.0 40 Trachysida a. aspera (LeConte) 0.3 0.2 0.3 0.0 0.0 0.3 1.0 3.0 0.0 0.4 16 Tragosoma depsarium (Linneaus) 0.0 0.1 0.3 0.1 0.3 0.0 0.0 0.0 0.0 0.2 8 Xestoleptura tibialis (LeConte) 0.0 0.2 0.8 0.2 0.2 0.0 1.0 0.0 0.4 0.0 17 Xylotrechus longitarsis {Casey)/undatulus (Say) 0.0 0.5 1.1 0.6 0.8 0.1 0.0 1.0 1.2 0.4 38 Cerylonidae Cerylon castaneum Say 0.6 0.8 1.3 0.9 0.7 0.9 2.0 2.0 0.8 0.4 58 Chrysomelidae sp. # 1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Altica tombacina (Mannerheim) 0.0 0.2 0.6 0.3 0.2 0.0 0.0 0.0 0.0 1.2 19 Bromius obscurus (Linneaus) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Crepidodera sp. 0.0 0.0 0.2 0.2 0.2 0.0 0.0 0.0 0.0 0.2 6 Hippuriphila sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Orsodacne sp. 0.0 0.0 0.0 0.0 0.2 0.1 0.0 0.0 0.4 0.0 4 Orsodacne atra (Ahrens) 0.1 0.7 0.2 0.5 0.3 0.6 0.0 0.0 4.2 0.4 45 Pachybrachis melanostictus Suffrain 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.2 0.0 2 Phaedon laevigatas (Duftschmid) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Phyllotreta striolata (Fabricus) 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Plateumaris rufa (Say) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Syneta pilosa W.J. Brown 0.0 0.6 0.4 0.0 0.0 0.3 0.0 0.0 0.0 0.0 13 Syneta albida LeConte 0.3 1.3 1.3 0.0 0.2 0.6 0.0 1.0 0.6 0.0 39 Syneta hamata Horn 0.0 0.1 0.0 0.0 0.7 0.1 0.0 0.0 0.0 0.0 6 Tricholochmaea sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Ciidae sp. # 1 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 sp.#2 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.2 0.0 3 sp.#3 0.0 0.3 0.0 0.1 0.2 0.0 0.0 0.0 0.2 0.0 6 Cis angustus Hatch 0.0 0.2 0.6 0.2 0.3 0.0 0.0 0.0 0.0 0.2 14 Cis sp. (fuscipes) Mellie 0.3 0.3 0.0 0.3 0.0 0.0 0.0 0.0 0.2 0.0 10 Diphyllcis ? sp. 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Dolichocis manitoba Dury 0.1 0.7 0.4 0.2 0.2 0.0 0.0 1.0 0.0 0.2 18 Octotemnus denudatus Casey 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 0.0 2 Orthocis punctatus Casey 1.4 0.4 0.5 0.3 0.2 0.3 0.0 2.0 0.2 0.0 32 Plesiocis sp. 0.0 0.3 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4 Xestocis sp. 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Calyptomerus oblongulus Mannerheim 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Cleridae Enoclerus sphegeus (Fabricus) 0.2 1.0 0.5 0.0 0.0 0.0 0.0 0.0 0.2 0.0 19 Enoclerus nr. Scheaferi Barr 0.1 0.4 0.0 0.0 0.0 0.0 3.0 0.0 0.0 0.0 8 Thanasimus undatulus (Say) 120 116 75.8 29.4 28.5 2.0 44.0 5.0 0.6 0.4 3850 Trichodes ornatus hartwegianus A. White 0.0 0.0 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.4 5 Coccinellidae Adalia bipunctata 0.0 0.0 0.2 0.8 0.3 0.0 0.0 0.0 0.0 0.2 13

83 (Linneaus) Coccinella septumpunctata Linneaus 0.0 1.0 2.3 3.7 2.3 0.0 0.0 4.0 5.8 3.2 136 Coccinella trifasciata perplexa Mulsant 0.0 0.1 0.5 0.8 1.5 0.0 1.0 1.0 0.8 2.0 39 Didion punctatum (Melsheimer) 0.0 0.1 0.3 0.2 0.2 0.0 0.0 0.0 0.0 0.4 9 Hippodamia tredecimpunctata (Say) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Macronaemia episcopalis (Kirby) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Mulsantina picta (Randall) 0.2 0.3 0.2 0.1 0.2 0.0 0.0 0.0 0.0 0.0 9 Psyllobora vigintimaculata (Say) 0.0 0.2 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 4 Scymnus sp. 0.0 0.0 0.1 0.0 0.3 0.0 0.0 0.0 0.0 0.0 3 Colydiidae Lasconotus complex LeConte 0.0 0.2 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 3 Lasconotus intricatus Kraus 0.1 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Corylophidae Molamba obesa Casey 0.0 0.2 0.0 0.0 0.0 0.1 1.0 0.0 0.0 0.6 7 Sacium lugubre LeConte 0.0 4.5 1.3 1.6 0.0 0.1 1.0 0.0 0.4 0.4 85 Antherophagus sp. # 1 0.4 0.2 0.4 0.1 0.2 0.0 4.0 0.0 0.0 0.0 16 Antherophagus sp. # 2 0.1 0.5 0.8 0.1 0.2 0.0 0.0 0.0 0.4 0.0 20 Atomaria sp. # 3 0.0 0.2 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Atomaria sp. # 4 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Atomaria sp. # 6 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Atomaria sp. # 9 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Atomaria sp. # 10 0.0 0.0 0.0 0.3 0.0 0.0 0.0 1.0 0.0 0.0 4 Atomaria sp. # 11 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Atomaria sp.#12 0.1 0.4 0.1 0.1 0.0 0.0 0.0 1.0 0.2 0.0 9 Atomairia sp. #13 0.0 0.0 0.2 0.0 0.2 0.0 0.0 0.0 0.0 •0.0 3 Atomaria sp. # 15 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Caenocelis sp. # 1 0.3 2.3 1.6 0.6 0.5 0.3 0.0 2.0 0.6 1.0 67 Cryptophagus sp. # 1 0.4 0.4 0.1 0.0 0.3 0.3 0.0 0.0 0.0 0.0 13 Cryptophagus sp. # 2 0.4 0.2 0.2 0.2 0.2 0.0 0.0 0.0 0.2 0.0 12 Cryptophagus sp. # 3 0.3 0.3 1.0 1.1 1.5 0.7 0.0 0.0 0.8 0.8 50 Cryptophagus sp. # 4 0.7 0.6 0.8 0.5 0.2 2.1 1.0 2.0 0.6 0.8 54 Cryptophagus sp. # 5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Henoticus sp. # 1 0.1 0.9 0.4 0.0 0.0 0.0 1.0 0.0 0.0 0.0 16 Henotiderus lorna (Hatch) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Myrmedophila americana (LeConte) 0.0 0.4 0.5 0.2 0.3 0.0 0.0 1.0 0.2 0.8 19 Salebius nr. minax 0.4 0.0 0.2 0.0 0.0 0.6 0.0 0.0 0.0 0.0 10 Cucujidae Cucujus claviceps Mannerheim 4.0 4.0 1.3 0.4 1.0 1.9 3.0 6.0 0.2 0.0 131 Dendrophagus cygnaei Mannerheim 1.7 5.0 1.5 0.5 0.3 0.3 0.0 2.0 0.2 1.0 106 Laemophloeus biguttatus (Say) 0.0 0.0 0.0 0.2 1.5 0.0 0.0 0.0 0.0 0.2 12 (Herbst) 0.0 0.3 0.4 0.2 0.2 0.0 0.0 0.0 0.2 0.2 12 Pediacus fuscus Erichson 0.2 4.6 9.2 1-4 1.5 0.0 4.0 4.0 1.8 0.8 198 Curculionidae Carphonotus testaceus Casey 0.3 0.1 0.1 0.0 0.3 0.0 0.0 0.0 0.2 0.2 9 Ceutorhynchus erysimi (Fabricius) 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Ceutorhynchus punctiger Gyllenhal 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.2 3 Cossonus pacificus Van Dyke 0.2 1.1 0.6 0.5 0.2 0.0 2.0 0.0 0.2 0.0 30

84 Magdalis alutacea LeConte 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Phloeophagus canadensis Van Dyke 0.0 0.0 0.2 0.0 0.0 0.0 0.0. 0.0 0.0 0.0 2 Pissodes fasciatus LeConte 0.2 1.6 0.5 0.0 0.0 0.7 3.0 0.0 0.4 0.2 37 Pissodes (nr.) fiskei Hopkins 0.0 0.2 0.0 0.0 0.0 0.0 1.0 0.0 0.0 0.0 3 Pissodes striatulus (Fabricus) 0.0 1.3 0.3 0.0 0.0 0.1 7.0 0.0 0.0 0.0 25 Pissodes striatulus dubius Randall 0.0 0.5 0.0 0.0 0.0 0.0 1.0 0.0 0.0 0.0 8 Proctorus decipiens (LeConte) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Rhyncolus brunneus Mannerheim 0.2 0.2 0.5 0.2 0.3 0.0 0.0 0.0 0.0 0.2 14 Rhyncolus macrops Buchanan 1.7 2.6 2.5 0.9 0.8 1.6 3.0 3.0 0.2 1.2 111 Sitona cylindricollis (Fahraeus) 0.0 0.0 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 2 Sitona lineellus (Bonsdorff) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 0.0 2 Tychius picirostris (Fabricus) 0.0 0.0 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 2 Dermestidae Anthrenus pimpinellae Fabricus 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Dermestes sp. 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Dermestes lardarius Linneaus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Dermestes talpinus Mannerheim 0.1 0.2 0.2 0.0 0.0 0.1 0.0 0.0 0.2 0.0 7 Megatoma sp. (cylindrica) (Kirby) 0.6 3.7 0.9 0.7 0.0 0.9 2.0 3.0 0.0 0.0 75 Megatoma verigatta (Horn) 1.1 2.1 1.8 0.8 0.2 0.7 13.0 0.0 0.2 0.2 84 Orphilis subnitidus LeConte 0.0 0.4 0.2 0.1 0.2 0.0 0.0 0.0 0.0 0.0 8 Pseudohadrotoma sp. (perversa) (Fall) 0.0 0.5 0.5 0.1 0.0 0.0 1.0 0.0 0.0 0.0 12 Laricobius laticollis Fall 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Dytiscidae sp. # 621 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Dytiscidae sp. # 623 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. # 624 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 sp. # 625 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. # 626 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Agabus sp. # 619 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Agabus sp. # 622 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Hydaticus aruspex Clark 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Hydroporus sp. # 1 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Hydroporus sp. # 2 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 2 Hydroporus sp. # 3 0.0 0.1 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 4 Hygrotus impressopunctatus (Schaller) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Rhantus binotatus (Harris) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Rhantus frontalis (Marsham) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Elateridae Agriotella occidentalis W.J. Brown 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.4 0.2 3 Ampedus behrensi + phelpsi (Horn) 0.0 2.9 14.5 4.2 7.2 0.0 2.0 2.0 1.01 4.2 301 Ampedus brevis (Van Dyke) 0.7 3.5 5.5 4.3 4.0 2.4 6.0 5.0 3.4 3.6 235 Ampedus mixtus (miniipennis?) (Herbst) 0.0 0.9 1.6 0.8 1.2 0.3 3.0 0.0 0.4 0.6 53 Ampedus moerens (LeConte) 0.0 1.8 1.2 1.1 1.2 0.0 0.0 0.0 0.4 0.6 56

85 Ampedus (nr.) moerens (LeConte) 0.0 4.0 8.4 1.7 3.7 0.0 1.0 0.0 0.2 4.6 200 Ampedus nigrinus (Herbst) 0.6 14.5 23.1 14.0 10.3 0.1 30.0 20.0 2.8 12.8 750 Ampedus occidentalis Lane 0.2 3.5 5.6 3.7 3.8 0.0 25.0 21.0 4.6 4.8 254 Ampedus phoenicopterus Germar 0.0 0.0 0.2 0.1 0.2 0.0 1.0 0.0 0.8 0.0 9 Ampedus pullus Germar 0.0 3.2 3.0 5.0 5.5 0.1 2.0 1.0 4.0 6.0 205 Athous nigropilis Motschulsky 0.4 . 0.0 0.1 0.0 0.0 0.3 0.0 0.0 0.0 0.0 7 Athous rufiventris rufiventris (Eschscholtz) 0.9 1.5 7.1 1.0 1.8 2.1 4.0 3.0 2.4 2.6 167 Cardiophorus (prob) tenebrosus LeConte 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Ctenicera sp. 134 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Ctenicera aeripennis (Kirby) 0.2 3.5 20.0 24.8 19.3 0.0 6.0 12.0 4.2 14.4 736 Ctenicera angusticollis (Mannerheim) 0.5 0.9 0.6 1.0 1.2 0.9 0.0 0.0 0.4 1.2 54 Ctenicera bipunctata (W.J. Brown) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Ctenicera bombycina (Germar) 0.0 0.3 0.4 0.6 0.0 0.1 0.0 0.0 1.0 0.2 20 Ctenicera crestonensis (W.J. Brown) 0.0 0.5 0.0 0.3 0.3 0.0 0.0 0.0 0.0 0.0 10 Ctenicera comes (W.J. Brown) 0.0 0.0 0.0 0.1 0.0 0.1 0.0 1.0 0.0 0.0 3 Ctenicera hoppingi (Van Dyke) 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Ctenicera kendalli Kirby 0.0 0.2 0.4 0.1 0.7 0.0 0.0 0.0 0.2 0.4 14 Ctenicera lobata (Eschscholtz) 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.4 0.0 4 Ctenicera lutescens (Fall) /sagitticollis (Eschscholtz) 0.2 1.5 0.6 0.5 0.2 0.3 1.0 0.0 0.2 0.0 35 Ctenicera mendax (LeConte) 0.0 0.0 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.2 4 Ctenicera nebraskensis (Bland) 0.4 5.6 8.8 9.6 5.2 1.4 1.0 0.0 5.2 24.6 452 Ctenicera nigricollis (Bland) 1.5 2.5 3.7 3.5 6.7 1.7 8.0 0.0 1.8 10.2 238 Ctenicera nitidula (LeConte) 0.0 0.1 0.4 0.3 0.0 0.0 2.0 0.0 0.6 0.2 14 Ctenicera pudica (W.J. Brown)+propo/a columbiana (Leconte) 5.5 9.3 10.4 27.2 45.7 5.1 43.0 1.0 20.2 28.6 1146 Ctenicera r. resplendens (Eschscholtz) 0.2 3.8 11.6 18.6 15.8 0.0 4.0 4.0 9.0 13.8 583 Ctenicera semimetallica (Walker) 0.0 0.2 0.4 0.3 0.7 0.1 0.0 0.0 0.4 1.0 21 Ctenicera triundulata (Randall) 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Ctenicera umbricola (Eschscholtz) 0.7 11.6 18.8 14.8 50.8 0.4 4.0 6.0 24.6 33.8 1101 Ctenicera volitans (Eschscholtz) 2.6 1.9 1.0 2.3 1.3 0.4 4.0 1.0 2.2 0.8 112 Dalopius (nr.) tristis W.J. Brown 0.0 0.5 1.5 1.0 1.7 0.0 0.0 0.0 1.2 2.6 61 Danosoma brevicorne (LeConte) 0.0 1.7 3.3 5.6 3.8 0.0 2.0 0.0 3.2 5.6 180 Drasterius debilis LeConte 0.5 1.1 0.2 0.5 0.0 3.0 4.0 1.0 0.6 0.4 55 Eanus sp. # 1 0.0 0.0 0.0 0.0 0.0 0.6 1.0 0.0 0.2 0.0 6 Eanus decoratus (Mannerheim) 0.0 0.0 0.1 0.0 0.0 0.0 1.0 0.0 0.2 0.0 3 Hypnoidus bicolor (Eschscholtz) 0.0 0.5 0.7 0.2 0.5 0.0 0.0 1.0 1.6 0.2 29 Hypnoides impressicollis (Mannerheim) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Lacon rorulentus (LeConte) 0.0 2-4 0.9 0.4 2.0 0.0 1.0 2.0 0.0 0.2 56

86 Limonius aeger LeConte 0.0 0.3 0.3 0.2 0.8 0.0 1.0 0.0 0.2 0.0 15 Limonius pectoralis LeConte 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.2 0.2 6 Negastrius tumescens LeConte 1.0 29.1 10.6 3.4 5.8 0.0 56.0 10.0 1.0 3.6 605 Neohypdonas tumescens (LeConte) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 brunneus 0.0 0.0 0.1 0.4 1.3 0.0 1.0 0.0 0.0 0.0 18 Erotylidae Triplax californica LeConte 0.4 1.0 0.9 1.8 1.3 0.7 1.0 0.0 0.8 1.2 67 Triplax antica LeConte 0.0 0.1 0.0 0.0 0.2 0.1 0.0 0.0 0.2 0.0 4 Triplax dissimulator (Crotch) 0.1 0.1 0.0 0.0 0.2 0.1 0.0 0.0 0.0 0.0 4 Eucnemidae Epiphanis sp. # 338 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Epiphanis cornutus Eschscholtz 0.2 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.2 4 Eucinetidae Eucinetus (nr.) oviformis LeConte . 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Histeridae Cylistus coarctatus (LeConte) 0.0 0.6 0.7 0.0 0.0 0.0 1.0 1.0 0.0 0.0 17 Gnathoncus barbatus Bosquet & Laplante 0.0 0.5 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 7 Gnathoncus communis (Marseul) 0.0 0.0' 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Margarinbtus rectus (Casey) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 .0.0 0.0 1 Paromalus mancus Casey 0.6 4.3 0.9 0.1 0.2 0.0 4.0 6.0 0.0 0.2 76 Platysoma coarctatum LeConte 0.0 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Platysoma leconti Marseul 0.0 0.1 0.5 0.5 0.0 0.0 0.0 0.0 0.2 0.0 12 Plegaderus setulosus Ross 0.0 0.1 0.0 0.0 0.0 0,0 0.0 0.0 0.0 0.0 1 Plegaderus sayi Marseul 0.0 0.1 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 3 Saprinus lugens Erichson 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Teretruis montanus Horn 0.0 0.2 0.4 1.1 2.2 0.0 0.0 0.0 0.8 0.2 35 Hydraena sp. (pacifica) Perkins 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 4 Ochthebius sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Hydrophilidae Cerycon sp. # 1 (herceus frigidus) Smetana 0.0 0.2 0.1 0.6 0.0 0.0 0.0 0.0 0.2 0.0 10 Cercyon sp. # 2 (cinctus) Smetana 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.4 2 Cercyon sp. # 3 (tolfino) Hatch 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Enochrus hamiltoni (Horn) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 orientalis Motscholsky /sempervarians Angus 0.0 0.4 0.4 0.2 0.3 0.0 0.0 0.0 0.0 0.0 12 Hydrobius fuscipes (Linneaus) 0.0 0.3 0.3 0.2 0.0 0.0 0.0 0.0 0.6 0.2 12 Laccobius sp. 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Laccobius borealis Cheary /carri D.C. Miller 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.0 0.4 0.4 8 Sphaeridium bipustulatum Fabricius 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Sphaeridium lunatum Fabricius 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Lampyridae Ellychnia corrusca (Linneaus) 0.0 0.0 0.3 0.0 0.0 0.0 o:o 1.0 0.2 0.0 5 Phausis rhombica Fender 0.0 0.0 0.0 0.1 0.2 0.1 0.0 0.0 0.2 0.0 4 Lathridiidae Corticarina (prob) cavicollis (Mannerheim) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Corticaria n. sp. 4.3 5.4 5.5 1.9 0.5 2.6 1.0 1.0 1.0 1.8 218 Corticaria sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1

87 Enicmus mendax Fall 0.1 0.1 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 4 Enicmus tenuicornis LeConte 2.6 2.5 2.1 1.2 1.3 2.7 0.0 3.0 1.8 1.0 132 Lathridius n. sp. 0.5 0.2 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 7 Lathridius ventralis 0.0 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.2 0.0 4 Stephostethus breviclavus (Fall) 0.4 0.4 0.2 0.3 0.0 0.0 0.0 0.0 0.0 0.0 13 Stephostethus cinnamopterus (Mannerheim) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Stephostethus liratus (LeConte) 0.9 0.5 0.2 0.5 0.2 0.4 0.0 0.0 1.0 0.0 30 Leiodidae Agathidium spp. 0.3 0.8 1.0 0.7 1.5 0.4 0.0 0.0 0.6 0.2 46 Agathidium sp. # 1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Agathidium basalis 0.0 0.0 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 2 Agathidium difformis (LeConte) 0.0 0.0 0.2 0.1 0.0 0.3 1.0 0.0 0.0 0.0 7 Agathidium depressum Fall /obtusum Hatch 3.7 3.5 3.1 0.9 1.0 2.7 2.0 6.0 1.0 1.2 163 Anisotoma globososa Hatch 0.0 0.5 1.2 1.2 0.2 0.4 0.0 1.0 1.2 0.4 43 Colon sp. # 1 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.2 0.0 2 Colon asperatum Horn 0.0 0.4 0.4 0.4 0.3 0.0 0.0 0.0 0.2 0.6 18 Colon (mylochus) aedeagosum Hatch 0.0 0.1 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 2 Colon magnicolle Mannerheim 1.2 1.4 1.5 0.6 0.7 0.0 0.0 1.0 0.0 0.4 57 Cyrtusa luggeri Hatch 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.4 0.2 4 Cyrtusa sp. (subtestacea) (Gyllenhal) 0.0 0.0 0.1 0.0 0.3 0.0 0.0 0.0 0.0 0.0 3 Hydnobius sp. # 1 0.0 0.5 0.7 0.2 1.5 0.0 0.0 0.0 0.8 1.0 34 Hydnobius sp. # 2 0.0 0.2 0.3 0.2 0.7 0.0 0.0 0.0 0.4 0.8 17 Hydnobius sp. # 3 0.0 0.1 0.0 0.2 0.0 0.1 0.0 0.0 0.2 0.2 6 Hydnobius pumilus LeConte 0.0 0.5 0.9 0.7 0.3 0.0 0.0 1.0 0.2 1.0 32 Leoides sp. # 49 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Leoides collaris (LeConte) 0.0 0.1 0.0 0.1 0.2 0.0 0.0 0.0 0.2 0.4 7 Leoides rufipes (Gebler) 0.0 0.0 0.2 0.2 0.0 0.1 1.0 1.0 0.0 0.8 11 Leoides sp. # 3 0.0 0.0 0.1 0.2 0.2 0.0 0.0 0.0 0.0 0.2 5 Leoides puncticollis C.G. Thomson / curvata /Mannerheim 0.0 0.2 0.3 0.3 0.5 0.0 0.0 1.0 0.6 0.2 16 Leiodes strigata (LeConte) 0.0 4.8 1.4 0.5 0.7 0.0 1.0 0.0 0.2 0.2 80 Triarthron lecontei Horn 0.0 0.1 0.1 0.0 0.0 0.1 1.0 0.0 0.0 0.2 5 Lucanidae Platycerus marginalis Casey 0.0 0.1 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.8 13 Lycidae Dyctyopterus spp. 2.3 0.7 2.4 1.2 0.7 1.0 2.0 6.0 0.4 0.8 94 Melandryidae Canifa sp. # 1 0.0 0.0 0.2 0.3 0.0 0.0 0.0 0.0 0.2 0.0 6 Emmesa stacesmithi Hatch 0.4 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6 Hallomenus sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Melandrya striata Say 0.1 0.1 0.4 0.2 0.3 0.0 0.0 0.0 0.0 0.0 10 Phryganophilus collaris LeConte 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Scotochroa basalis LeConte 0.1 0.4 0.7 0.5 0.0 0.6 1.0 0.0 0.6 0.2 27 Serralopalpus substriatus Haldeman 0.6 1.5 0.7 0.1 0.0 0.6 2.0 0.0 0.0 0.0 38 Xyleta laevigata (Hellenius) 2.9 87.7 27.1 8.4 6.0 0.4 82.0 70.0 6.2 7.4 1635 Zilora occidentalis Mank 0.2 0.0 0.1 0.0 0.2 0.1 0.0 0.0 0.0 0.0 5 Melyridae Attalus sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Hoppingiana sp. 0.2 0.2 1.1 0.3 0.0 0.0 0.0 0.0 0.4 0.2 23

88 (hudsonica) (LeConte) Semijulistus ater (LeConte) 0.0 3.3 3.2 3.6 2.3 0.0 0.0 2.0 2.6 2.0 146 Trichochrous albertensis Blaisdell 0.4 0.2 0.5 0.5 0.7 2.9 0.0 0.0 0.0 0.4 43 Mordellidae Tomoxia borealis (LeConte) 0.0 0.1 0.3 0.2 0.2 0.0 0.0 0.0 0.2 0.2 9 Mycetophagidae Mycetophagus distinctus Hatch 0.0 0.6 0.5 0.3 0.3 0.4 0.0 0.0 1.0 0.4 27 Mycetophagus tenuifasciatus Horn 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 (Linneaus) 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 (prob) turbans Kuschel 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Pityomacer pix Kuschel 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Nitidulidae #296 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 1 #299 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 .0.0 2 # 339 + # 357 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Colopterus truncatus (Randall) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Epuraea sp. # 1 0.2 3.1 0.9 0.3 0.2 0.3 2.0 0.0 0.8 0.2 59 Eupuraea sp. # 2 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 sp.#3 0.0 0.1 0.1 0.1 0.0 0.0 1.0 0.0 0.0 0.0 4 sp.#4 0.0 0.2 0.0 0.0 0.2 0.0 0.0 0.0 0.2 0.0 4 sp.#6 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 sp.#7 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 1 sp.#8 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. #9 0.0 0.4 0.1 0.0 0.3 0.0 0.0 0.0 0.0 0.0 7 sp. # 10 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 sp. # 11 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 sp. # 12 0.0 0.3 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4 sp. # 13 0.0 0.5 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6 sp. # 14 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. # 15 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4 Epuraea depressa 1.1 0.1 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 15 Epuraea flavomaculata Maklin 0.1 0.1 0.0 0.0 0.7 0.0 0.0 0.0 0.4 0.4 10 Epuraea planulata Erichson 0.1 1.5 1.5 0.3 0.2 0.3 3.0 3.0 0.2 0.4 48 Epuraea (nr.) populi Dodge 0.0 0.2 0.1 0.0 0.2 0.0 0.0 0.0 0.0 0.0 4 Eupraea terminalis Mannerheim 0.2 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 3 Eupraea truncatella Mannerheim 0.3 2.1 1-4. 0.1 0.2 0.0 2.0 0.0 0.2 0.0 46 Glischrochilus confluentus (Say) 0.1 2.6 0.2 0.1 0.2 0.0 0.0 0.0 0.0 0.2 35 Glischrochilus moratus W.J. Brown 0.0 0.2 0.1 0.0 0.3 0.0 0.0 0.0 0.0 0.2 6 Glischrochilus quadrisignatus (Say) 0.0 0.4 0.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 9 Omosita discoidea (Fabricius) 3.5 1.3 0.6 0.1 0.0 0.3 2.0 0.0 0.0 0.0 59 Thalycra mixta H. Howden 0.0 0.4 0.5 0.0 0.0 0.4 0.0 0.0 0.2 0.0 14 Oedemerinae Calopus angustus LeConte 1.0 1.5 3.0 1.0 0.5 0.7 2.0 0.0 0.0 1.4 87 Phalacridae sp. #1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 Pselaphidae Pselaphus bellax Casey 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Ptinus californicus Pic 0.0 0.2 0.1 0.2 0.2 0.0 0.0 0.0 0.2 0.2 8 Pyrochroidae Dendroides ephemeroides (Mannerheim) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Pythidae Priognathus monilicornis LeConte 0.0 1.1 0.3 0.1 0.5 0.0 0.0 0.0 0.0 0.0 19

89 Pytho sp. # 1 0.0 0.1 0.3 0.2 0.2 0.0 1.0 1.0 0.2 0.0 10 Pytho sp. # 2 0.3 0.1 0.1 0.0 0.0 0.1 1.0 0.0 0.0 0.0 7 Rhizophagidae Rhizophagus pseudobrunneus Bousquet 0.2 0.2 0.2 0.1 0.0 0.0 0.0 0.0 0.4 0.0 9 Rhyzophagus dimidiatus Mannerheim 2.6 0.5 0.1 0.2 0.2 0.1 1.0 1.0 0.0 0.0 38 Rhizophagus remotus LeConte 0.6 0.8 0.7 0.1 0.5 0.6 0.0 0.0 0.4 0.2 34 Salpingidae Rhinosimus viridiaeneus Randall 2.1 1.8 0.7 1.0 1.0 0.1 0.0 0.0 0.0 0.4 68 alternatus (LeConte) 0.0 0.4 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0 7 Sphaeriestes sp. 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Scaphidiidae Scaphisoma castaneum Motschulsky 0.1 0.4 0.3 0.9 0.7 0.0 0.0 0.0 0.2 0.2 23 Scaphium sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Scarabaeidae Aegialia rufescens Horn 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. # 328 0.0 0.0 0.0 0.0 0.0 0.1 2.0 1.0 0.0 0.0 4 sp. # 608 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 sp. # 609 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 sp. # 610 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 sp.#611 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.2 0.2 3 Aphodius distinctus (O.F. Muller) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Aphodius fimetarius (Linneaus) 0.8 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 10 Aphodius haemorrhoidalis (Unneausypectoralis LeConte 0.1 0.1 0.2 0.1 1.0 0.0 0.0 0.0 0.0 0.2 12 Aphodius leopardus Horn 0.0 0.0 0.3 0.0 0.2 0.7 0.0 0.0 0.0 0.0 9 Aphodius opacus LeConte 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Dichelonyx vicina (Fall) 0.0 0.0 0.0 0.0 0.5 0.0 0.0 0.0 0.0 0.2 4 Diplotaxis brevicollis LeConte 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 2 Scirtidae Cyphon sp.(p) 0.1 0.3 1.1 1.5 0.7 0.0 0.0 0.0 2.4 1.0 52 Cyphon concinnus (LeConte) 0.0 0.0 0.2 0.0 0.2 0.0 0.0 0.0 0.0 0.4 5 Scolytidae Carphoborus vandykei Bruck 0.0 0.5 0.3 0.3 0.0 0.0 0.0 0.0 0.2 0.0 13 Cryphalus ruficollis Hopkins 0.1 0.7 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.2 13 Dendroctonus pseudotsugae Hopkins 7699 1285910362 4152 1137 6.4 16.0 15.0 7.6 2.0 380785 Dendroctonus rufipennis (Kirby) 0.0 0.1 0.2 0.1 0.0 0.6 0.0 1.0 0.4 0.0 11 Dryocetes affaber (Mannerheim) 0.5 7.4 1.4 0.8 0.2 0.6 6.0 4.0 0.8 0.4 130 Dryocetes autographus (Ratzeburg) 1.1 18.8 5.9 2.1 0.7 0.6 14.0 4.0 2.0 2.4 352 Dryocetes betulae Hopkins 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Dryocetes caryi Hopkins /schelti Swaine 0.2 0.2 0.0 0.0 0.0 0.0 0.0 1.0 0.0 0.0 5 Dryocetes confusus Swaine 0.3 4.1 0.3 0.0 0.0 0.1 1.0 0.0 0.0 0.0 53 Gnathotrichus retusus LeConte 2.2 15.7 30.7 12.7 0.7 0.6 4.0 2.0 4.4 1.4 703 Hylastes nigrinus (Mannerheim) 13.2 140 120 44.2 8.2 6.1 224 13.0 11.4 13.2 3903 Hylastes longicollis Swaine 0.3 0.8 0.4 0.8 0.3 0.1 1.0 0.0 0.0 0.4 30 Hylastes ruber Swaine 5.1 54.2 34.5 9.0 10.3 7.3 26.0 1.0 3.2 7.0 1307 Hylurgops porosus (LeConte) 0.0 105 20.4 3.5 1.7 0.1 82.0 2.0 0.2 0.6 1522 Hylurgops reticulatus Wood 0.0 0.2 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3

90 Hylurgops rugipennis Mannerhiem 0.0 0.8 0.0 0.1 0.0 0.1 1.0 0.0 0.0 0.0 12 Ips latidens (LeConte) 0.0 0.2 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 4 Ips mexicanus (Hopkins) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Ips perturbatus (Eichhoff) 0.1 5.3 0.8 0.0 0.2 0.0 4.0 2.0 0.6 0.2 79 Ips pini (Say) 0.1 1.7 0.5 0.0 0.5 0.0 5.0 1.0 0.0 0.2 35 Ips tridens (Mannerheim) 0.0 3.4 0.9 0.1 0.3 0.0 2.0 1.0 0.0 0.2 54 Orthotomicus caelatus (Eichhoff) 0.1 4.0 5.7 1.5 0.2 0.0 10.0 2.0 0.6 0.2 140 Phloeosinus pini Swaine 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Phloeotribus lecontei Schedl/p/'cea Swaine 0.0 1.5 0.3 0.2 0.0 0.1 0.0 0.0 0.0 0.0 23 Pityogenes hopkinsi Swaine 0.1 0.1 0.4 0.1 0.2 0.0 0.0 0.0 0.0 0.0 8 Pityogenes knetchteli Swaine 0.0 0.7 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 9 Pityogenes plagiatus (LeConte) 0.1 4.2 1.0 0.3 0.0 0.0 2.0 0.0 0.2 0.2 65 Pityokteines elegans Swaine 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Pityokeines minutus (Swaine) 0.0 0.5 0.5 0.5 0.0 0.0 0.0 0.0 0.0 0.0 15 Pityopthorus sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Pityophthorus nitidulus Swaine(+ tuberculatus Eschhoff) 0.1 0.8 2.2 0.9 0.5 0.3 2.0 1.0 1.4 0.4 60 Pityophthorus opaculus LeConte 0.0 0.6 0.3 0.4 0.2 0.0 0.0 0.0 0.6 0.0 18 Pityophthorus pseudotsugae Swaine 0.1 3.2 0.5 0.2 0.0 0.0 2.0 0.0 0.2 0.0 46 Pityopthorus aquilus Blackman (+ aplanatus) 0.0 0.2 0.1 0.1 0.2 0.0 2.0 0.0 0.0 0.0 7 Polygraphus convexifrons Wood 0.4 1.0 0.2 0.1 0.0 0.1 0.0 0.0 0.2 0.0 20 Polygraphus rufipennis (Kirby) 3.2 43.2 21.1 7.8 4.3 0.3 12.0 6.0 5.8 2.2 903 Pseudohylesinus nebulosus LeConte 4.3 102 61.5 3.9 2.3 4.7 647 0.0 1.6 1.2 2596 Scierus annectans LeConte 14.9 9.6 4.3 0.6 0.8 4.4 12.0 2.0 0.8 0.0 362 Scierus pubescens Swaine 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Scolytus sp. (unispinosus) LeConte 0.0 15.7 14.2 1-2 0.2 0.1 6.0 0.0 1.4 0.8 360 Scolytus opacus Blackman 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Scolytus subscaber LeConte 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Scolytus piceae (Swaine) 0.0 4.7 0.6 0.3 0.3 0.0 0.0 0.0 0.2 0.2 66 Scolytus tsugae (Swaine) 1.3 9.2 1.0 0.1 0.0 0.0 7.0 0.0 0.0 0.0 133 Scolytus unispinosus LeConte 0.4 28.0 12.5 1.4 0.2 0.1 5.0 0.0 1.6 1.6 487 Trypodendron betulae Swaine 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 2 Trypodendron lineatum (Olivier) 1274 4013 485 3.4 1.3 13.0 598 2.0 1.0 0.2 62983 Trypodendron retusum (LeConte) 2.0 0.9 0.1 0.1 0.2 1.3 1.0 0.0 0.0 0.0 43 Trypodendron rufitarsis (Kirby) 0.7 0.0 0.1 0.0 0.0 0.3 0.0 0.0 0.0 0.0 10 Trypophloeus populi Hopkins 0.0 0.2 0.0 1.4 0.0 0.0 0.0 0.0 0.0 0.0 16 Xylechinus montanus Blackman 5.6 0.2 0.1 0.0 0.0 0.7 0.0 0.0 0.0 0.0 64 Scraptiidae Anaspis sp. 25.6 8.7 9.6 3.2 3.2 1.4 3.0 61.0 4.0 3.6 624 Anaspis sp. # 2 0.0 1.1 1.2 1.4 0.5 0.0 1.0 3.0 0.0 0.4 48 Hallomenus sp. 0.1 0.1 0.3 0.0 0.0 0.3 0.0 0.0 0.0 0.0 6

91 Orchesia (nr.) castanea (Melsheimer) 0.1 0.2 0.1 0.2 0.0 0.1 0.0 0.0 0.4 0.0 9 Orchesia ornate Horn 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Scraptiidae sp. # 3 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Scydmaenidae Stenichnus californicus Motschulsky 0.0 0.2 0.2 0.2 0.3 0.0 0.0 0.0 0.0 0.4 10 Silphidae Oiceoptoma noveboracense (Forster) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Thanatophilus lapponicus (Herbst) 0.0 0.4 0.2 0.2 0.0 0.0 0.0 1.0 0.0 0.0 9 Spaeritidae politus Duftschmid 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Sphindidae Odontosphindus clavicornis Casey 0.0 0.3 1.2 0.5 0.0 0.1 0.0 0.0 0.2 0.8 27 Staphylinidae Acidota crenata (Fabricius) 0.2 0.6 1.9 2.3 0.3 0.3 0.0 0.0 0.4 1.2 65 (misc.spp) 1.0 2.6 0.9 0.1 0.0 0.0 1.0 0.0 0.0 0.2 52 Aleochara castaneipennis Mannerheim 0.1 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 5 Aleochara gracilicornis Bernhauer 0.0 0.3 0.2 0.2 0.2 0.0 0.0 0.0 0.0 0.0 8 Aleochara (xeno) lanuginosa Gravenhorst 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Aleochara rubricalis Casey 0.0 0.1 0.5 0.1 0.0 0.0 0.0 0.0 0.0 0.0 8 Aleochara sekanai Klimaszewski 0.0 0.1 0.0 0.1 0.2 0.0 0.0 0.0 0.0 0.0 3 Aleochara suffosa (Casey) 0.0 0.9 0.4 0.1 0.2 0.0 0.0 0.0 0.2 0.0 17 Aleochara (aleo) tahoensis Casey 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 Aleochara villosa Mannerheim 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Amischa sp. 0.0 0.0 0.2 0.3 0.0 0.0 0.0 0.0 0.0 0.0 5 Anotylus rugosus (Farbricius) 0.0 0.5 0.2 0.4 0.0 0.0 0.0 0.0 0.0 0.0 11 Anotylus tetracarinatus (Block) 0.0 0.0 0.3 0.0 0.2 0.0 0.0 0.0 0.0 0.0 4 Anthobium reflexicolle Casey 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Atheta dentate Bernhauer 0.3 2.4 1.4 0.4 0.0 0.1 3.0 0.0 0.0 0.2 53 Atrecus macrocephalus (Nordmann) 0.6 0.3 0.3 0.2 0.0 0.0 0.0 0.0 0.0 0.0 14 Atrecus quadripennis (Casey) 0.1 0.0 0.0 0.1 0.0 0.0 1.0 0.0 0.0 0.0 3 Bledius ruficornis LeConte 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Bisnius picicornis (Horn) 0.3 1.7 2.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 46 Bolitopunctus muricatulus (Hatch) 0.9 1.0 2.0 1.2 2.0 3.0 1.0 0.0 2.4 2.0 111 Bryophacis spp. 0.0 0.1 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.2 6 Bryophacis arcticus 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Bryophacis Canadensis Campbell 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Bryophacis punctulatus (Hatch) 0.1 0.3 0.4 0.1 0.0 0.0 1.0 1.0 0.2 0.0 12 Bryophacis smetanai Smetana 0.0 0.1 1.0 0.5 1.0 0.0 0.0 0.0 0.4 0.4 27 Carphacis nepigonensis (Bernhauer) 0.4 1.2 1.5 0.3 0.5 0.1 1.0 2.0 0.0 0.4 45 Clavilispinus rufescens (Hatch) 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Creophilus maxillosus (Linneaus) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Dienopteroloma subcostatum (Maklin) 0.1 0.5 0.1 0.0 0.0 0.3 0.0 0.0 0.0 0.0 10 Earota sp. 1.0 1.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 23

92 Eucnecosum tenue (LeConte) 0.0 0.1 0.2 0.1 0.2 0.0 0.0 0.0 0.0 0.2 6 Eusphalerum spp. (mostly pothos (Mannerheim)) 202 2.9 2.7 1.1 3.7 6.9 6.0 1.0 3.6 2.2 2199 Gabrius picipennis (Maklin) 0.0 2.4 5.8 2.4 1.2 0.0 0.0 0.0 0.6 1.0 129 Gymnusa atra Casey 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Gymnusa sp. (grandiceps Casey) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Gymnusa pseudovariegata Klimaszewski 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.2 0.0 2 Gyrohypnus fracticornis (O.F. Muller) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Gyrophaena spp. 0.0 0.5 0.2 0.2 0.0 0.0 0.0 0.0 0.4 0.4 14 Grypeta sp. 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Hapalaraea sp. #1 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Hapalaraea dropephylla © 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Hapalaraea megarthroides (Fauvel) 0.1 1.0 0.0 0.1 0.0 0.4 2.0 0.0 0.0 0.0 18 Heterothops sp. 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Heterothops conformis Smetana 0.0 3.2 5.8 1.5 0.2 0.0 0.0 0.0 0.2 0.2 117 Heterothops fraternus Smetana 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Ischnosoma sp. (fimbriatum) Campbell 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Lathrobium negrum LeConte 0.0 0.3 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 4 Leptusa sp. 0.1 0.2 0.2 0.1 0.0 0.0 0.0 0.0 0.2 0.6 4 Linohesperus borealis (Casey) 0.0 0.2 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.4 10 Lordithon bimaculatus (Couper) 0.3 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 4 Lordithon cascadensis (Maklin) 0.0 0.0 0.0 0.2 0.3 0.1 0.0 0.0 0.6 0.6 11 Lordithon fungicola Campbell 1.2 0.4 0.2 0.0 0.0 0.0 0.0 1.0 0.2 0.4 22 Lordithon poecilus (Mannerheim) 0.0 0.1 0.2 0.0 0.2 0.0 0.0 0.0 0.0 0.0 4 Lordithon t. thoracicus (Fabricius) 0.0 0.1. 0.2 0.1 0.7 0.0 1.0 0.0 0.0 0.2 10 Myremcocephalus arizonicus (Casey) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Medon sp. (nr. Pallescens) 0.0 0.1 0.4 0.1 0.3 0.0 0.0 0.0 0.0 0.0 8 Megarthrus angulicollis Maklin 0.2 1.4 0.4 0.2 0.0 0.1 1.0 0.0 0.4 0.0 27 Micropeplus laticollis Maklin 0.1 0.0 0.3 0.1 0.2 0.1 0.0 0.0 0.2 0.0 8 Micropeplus smetanai Campbell 0.0 0.5 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0 8 Micetoporus sp. 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 Mycetoporus americanus Erichson 0.1 0.0 0.3 0.3 0.0 0.1 0.0 0.0 0.2 0.2 10 Mycetoporus brunneus (Marsham) 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.2 4 Mycetoporus rufohumoralis Campbell 0.4 1.0 0.5 0.4 0.5 1.9 1.0 0.0 0.4 0.6 47 Mycetoporus rugosus Hatch 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Myrmecocephalus arizonicus (Casey) 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.4 0.0 5 Neohypnus obscurus (Erichson) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 2 Nitidotachinus tachyporus 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Nudobius cephalus (Say) 0.0 3.1 1.8 0.9 0.3 0.0 0.0 1.0 0.2 0.6 71 Ochthephilus planus 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1

93 (LeConte) Olophrum boreale (Paykull) 0.0 0.1 0.1 0.2 0.0 0.0 0.0 0.0 0.4 0.0 6 Olophrum consimile (Gyllenhal) 0.0 0.0 0.0 0.2 0.0 0.0 0.0 1.0 0.6 0.6 9 Omalium sp. # 1 0.1 0.1 0.0 0.0 0.0 0.0 1.0 0.0 0.0 0.0 3 Omalium sp. # 2 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Omalium n. sp. 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Omalium spp. 0.0 0.3 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 4 Omalium sp. (foraminosum Maklin) 0.2 0.1 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0 6 Orus sp. 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Oxytelus sp. 0.0 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 8 Oxytelus fuscipennis Mannerheim 0.2 0.0 0.5 1.6 0.8 0.3 0.0 1.0 0.6 1.2 41 Pelecomalium testaceum (Mannerheim) 4.5 1.8 0.6 0.2 0.5 0.3 15.0 2.0 0.4 0.2 99 Philodrepa (?) Dropephylla sp. (nr. longula Maklin) 0.1 1.1 6.9 4.6 4.2 0.4 2.0 4.0 1.6 4.0 197 Philonthinii spp. 0.3 1.2 2.0 0.7 0.3 0.3 0.0 0.0 0.2 0.4 52 Philonthus couleensis Hatch 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Philonthus concinnus (Gravenhorst) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Philonthus crotch! Horn 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Philonthus furvus Nordmann 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 2 Philonthus politus (Linneaus) 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Philonthus varians (Paykull) 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Phloeopra sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Placusa tacomae Casey 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 Proteinus sp. 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Pseudopsis sp. 0.1 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 Quediini spp. 0.0 0.3 0.3 0.1 0.0 0.1 0.0 0.0 0.4 0.0 10 Quedius criddlei (Casey) 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4 Quedius (Disticalius) sp. 0.0 0.0 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 2 Quedius erythrogaster Mannerheim 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Quedius m. molochinoides Smetana 0.0 0.5 1.1 1.3 1.0 0.0 0.0 1.0 2.6 1.0 56 Quedius pediculus (Nordmann) 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Quedius plagiatus Mannerheim 3.5 0.7 0.6 0.4 0.2 1.6 3.0 0.0 0.2 0.0 70 Quedius rusticus/vilis Smetana 0.3 0.7 2.8 0.3 0.5 0.0 0.0 0.0 0.2 0.2 49 Quedius s. spelaeus Horn 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Quedius transparens Motschulsky 0.0 0.0 0.1 0.1 0.0 0.0 0.0 1.0 0.0 0.0 3 Quedius velox Smetana 4.4 7.3 8.9 5.3 4.8 4.7 9.0 16.0 2.6 4.8 399 Sepedophilus littoreus (Linneaus) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Siagonium stacesmithi Hatch 0.6 0.8 2.1 1.0 0.2 0.0 3.0 3.0 0.4 0.2 58 Sonoma parviceps (Maklin) 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 1 Staphylinus pleuralis LeConte 0.1 0.5 0.7 0.5 0.7 0.1 1.0 0.0 0.4 0.2 28 Stenus bilineatus J. Sahlberg 0.0 0.8 0.3 0.2 0.2 0.0 0.0 0.0 0.0 0.0 15 Stenus juno Paykull 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Stenus plicipennis (Casey) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1

91 Syntomium grahami Hatch 0.0 0.0 0.2 0.0 0.2 0.0 0.0 0.0 0.0 0.0 3 Tachinus basalis Erichson 0.7 0.1 0.1 0.1 0.3 0.7 0.0 0.0 0.2 0.4 20 Tachinus elongatus Gyllenhal 0.1 0.5 0.0 0.0 0.0 0.3 2.0 1.0 0.2 0.0 12 Tachinus frigidus Erichson 0.2 0.1 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.2 6 Tachinus nigricornis Mannerheim 0.1 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 2 Tachinus thruppi Hatch 0.1 0.0 0.1 0.2 0.0 0.0 0.0 0.0 0.2 0.2 6 Tachinus vergatus Campbell 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.0 1 Tachyporus sp. (canadensis CampbellJ 0.0 0.0 0.1 0.0 0.3 0.0 0.0 0.0 0.0 0.0 3 Tachyporus sp. (lecontei CampbellJ 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1 Tachyporus sp. 0.0 0.0 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 2 Trichophya pilicornis (Gyllenhal) 0.0 1.2 0.6 0.2 0.0 0.1 1.0 0.0 0.6 0.0 27 Zyras sp. 0.0 0.3 0.3 0.3 0.2 0.0 0.0 0.0 0.0 0.2 11 Stenotrachelus aeneus (Fabricius) 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Tenebrionidae Bius estriatus (LeConte) 0.0 1.5 0.3 0.0 0.0 0.3 0.0 0.0 0.2 0.0 23 Corticeus praetermissus (Fall) 0.1 0.3 0.2 0.0 0.0 0.0 1.0 0.0 0.0 0.2 8 Corticeus subopacus (Wallis) 0.2 0.2 0.4 0.1 0.0 0.1 2.0 0.0 0.0 0.0 12 Corticeus tenuis (LeConte) 0.0 0.1 0.0 0.1 0.0 0.0 0.0 0.0 0.4 0.0 4 Eleates explanatus Casey 0.3 0.2 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 6 Mycetochara fraterna (Say) 0.1 0.0 0.3 0.1 0.0 0.0 0.0 0.0 0.2 0.0 6 Phaleromela verigata Triplehorn 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Platydema sp. # 1 0.0 0.0 0.0 0.2 0.0 0.0 1.0 0.0 0.0 0.0 3 Platydema americanum Castelnau & Brulle 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Scaphidema aeneolum (LeConte) 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1 Tribolium audax Halstead 0.3 4.3 1.5 1.7 1.0 0.0 0.0 1.0 0.6 1.6 101 Upis ceramboides (Linneaus) 0.0 0.2 0.1 0.2 0.2 0.0 0.0 0.0 0.0 0.0 6 Tetratomidae Abstrulia (nr.) veriegatta Casey 0.9 0.6 0.4 0.3 0.3 1.6 3.0 0.0 0.0 0.0 39 Tetratoma concolor LeConte 1.9 0.2 0.3 0.0 0.0 1.4 0.0 0.0 0.0 0.2 35 Throscidae sp. # 573 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.0 0.0 1 Pactopus hornii (LeConte) 0.4 0.6 1.1 0.3 0.7 0.1 2.0 2.0 0.2 1.0 41 sp. (carnicollis (Schaeffer)J 0.0 0.1 0.0 0.1 0.2 0.0 0.0 0.0 0.0 0.0 3 Trixagus sp. 0.0 0.3 0.0 0.3 0.3 0.0 0.0 0.0 0.0 0.4 10 Trogostidae Calitys scabra (Thunberg) 0.4 1.3 2.0 4.7 4.3 0.4 0.0 5.0 1.0 1.8 135 Ostoma ferrugina (Linneaus) 0.6 0.1 0.4 0.6 0.2 0.3 1.0 1.0 0.4 0.4 26 Thymalus marginicollis Chevrolat 2.7 0.6 0.2 0.0 0.0 4.3 4.0 2.0 0.0 0.0 72

Mean trapped beetle abundance was significantly higher in pheromone-baited sites over control sites. Across all treatment years, on average, pheromone-

95 baited sites contained a 26-fold increase in the mean number of beetles/site over control treatments (9,943 vs 371 beetles; baited and control respectively). The greatest difference was observed in preharvest data (62 fold increase) and the least difference was observed in post 4/5 data (4.3 fold increase) (Figure 3.1).

100000

10000 -\

CO 1000 "55 o o a> 100 CO * c

O

Preharvest Post 1 Post 2 Post 3 Post4/5 Harvest Stage

Figure 3.1. Abundance of flying beetles in pheromone-baited (•) and unbaited* (•) Lindgren funnel traps trapped in preharvest through 475th season post harvest conditions. Habitat conditions were standing, or recently harvested, Douglas-fir

beetle attacked, mature Interior Douglas-fir (Fd8, >100 yrs), Fort St James Forest District, British Columbia. * Post 1 and post 2 unbaited data represent a single trapping site, and are included for general comparison only.

Whittaker plots (rank distribution) follow logarithmic species distributions for both baited and control data sets across all treatment levels/harvest years.

96 Despite variations in the length and the slope of data within and between treatments, in all cases pheromone-baited sites have a greater slope than

control sites. This slope difference is greatest for preharvest sites, lowest for

1st season postharvest sites (Figure 3.2).

The top ten abundant species for baited and control sites across all treatment

years is presented in Table 3.6. Of all species listed only the Scolytid

Hylastes nigrinus is found ranked in the top ten for all baited and control data

across all treatment years. Rank position of the species was variable both

within and between treatment years. The target species Douglas-fir beetle was inconsistently ranked in the top ten for 3/5 years of control data. In

pheromone-baited sites however, the beetle was the most abundantly trapped

species across all treatment years. Remaining species varied in frequency of

occurrence and rank. One trend observed in the ten most abundant species

is worth noting. In preharvest data, members of the family Scolytidae make

up 60% of the ten most abundant species (6/10 & 6/10 species: baited &

control sites respectively). This proportion increased in the first year after

harvesting (8/10 & 6/10 species: baited & control), but subsequently

decreased with every subsequent postharvest treatment year (3/10 & 1/10:

baited & control for the 415th postharvest year). As Scolytids decreased,

members of the family Elateridae increased, comprising at least half of the top

ten species in 4/5th season postharvest conditions (5/10 species & 7/10

species: baited & control sites respectively).

97 10000 100000

• Preharvest Baited 10000 Post-1 Baited 1000 Preharvest Control 1000 Post-1 Control

100 100

10

Species Rank (1-208) Species Rank (1-416) a. Preharvest b. First season postharvest'

100000

10000 -Post-3 Baited

1000 • Post-2 Control -Post -3 Control

100

10

1

0.1 —* •

0.01

Species Rank (1-413) Species Rank (1-350) c. Second season postharvest * d. Third season postharvest

10000

1000 i—Post-4/5 Control 100

10

1

Species Rank (1-262)

e. Fourth/fifth season postharvest

Figure 3.2. Species abundance of flying Coleoptera in a) preharvest habitat, b) 1st season postharvest habitat, c) 2nd season postharvest habitat, d) 3rd season postharvest habitat, and e) 4/5 season postharvest habitat. For all harvest years abundance is expressed by rank as mean abundance per site for pheromone- baited and unbaited/control sites. Beetles were trapped in Lindgren funnel traps in

mature Interior Douglas-fir (Fd8, >100 yrs) in the Fort St James Forest District, British Columbia. * Control data resulting from a single site included for general comparison only.

98 71 Q. CD CD 00 Ol 7? 3

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Nine measures of diversity were applied to each baited and control data set in each treatment year to assess changes in community structure (Figure 3.3): five measures of species richness (Number of species (S); Margalef (d),

Shannon-Wiener (H'i0), Brillouin, and Fisher (ex) indices), one measure of evenness (Pielou (J') index), one measure of dominance (Simpson index (1-

)), and two taxonomic based measures developed by Clarke and Warwick

(Taxonomic diversity (5), and taxonomic distinctness (8 *)).

Given the large volume of data contained in the analysis of diversity measures, the results have been broken down into two presentations: First, common and similar trends observed between indices are presented. This survey of trends is followed by assessments of individual measures of diversity, including trends and statistical differences between baited and control data, as well as between treatment years within an index.

Overview

Results revealed that between baited and control data, six out of nine measures were statistically different (at rx = 0.025) across all treatment years.

These results were observed in six out of 8 calculated indices including

Shannon-Wiener (H'IO), Brillouin, Fisher (

Number (S), and Margalef s index resulted in similar trends and values for

100 baited and control data. In contrast, the Taxonomic Distinctness index (8 *)) resulted in inconsistent values between baited and control data (Figure 3.3).

Across the range of diversity measures there were two reoccurring trends:

The first trend, observed in S and d richness indices, was a rapid increase in diversity from preharvest data to 1st / 2nd year postharvest data, followed by a gradual decrease in 3rd and 4/5th years. This trend occurred without significant difference for baited and control data in either index. The second trend

observed in richness indices H'i0, Brillouin, and a, was also observed in indices measuring dominance (1- ), evenness (J'), and taxonomic diversity

(8). It consisted of overall higher diversity for control traps, beginning with intermediate/high diversity in preharvest sites, followed by a drop in diversity associated with the single site data point for treatment year post 1. Diversity returned to or exceeded preharvest levels with the single site data point for treatment year post 2, and subsequently stabilized through post 3 and 4/5 data sets. The remaining two indices, a and 8 * followed unique trends described in their respective index assessments below.

Species number (S).

Mean species number (S) (Figure 3.3a) was lowest for baited and control data in preharvest sites (65.00, and 53.60 respectively) (Table 3.7). In pheromone- baited sites, richness peaked at 145.40 in the 2nd year postharvest, falling to

101 Post 1 Post 2 Post 3 Post 4/5 Harvest Year Harvest Year

g 0.3- % 0.2. te s Diversit y (Brillouin ; Spe c

Preharwst Post 1 Post 2 Post 3 Post 4/5 Preharvest Post 1 Post 2 Post 3 Post 4/5 Harvest Year Harvest Year

8 i 0.8

§—-§E S

Post 2 Post 3 Preharvest Post 1 Post 2 Post 3 Harvest Year Harvest Year

Preharvest Post 1 Post 2 Post 3 Post 415

Harvest Year

Figure 3.3. Measures of flying beetle diversity from pheromone-baited (•) and unbaited (•) funnel traps in Mature/Old

growth northern interior Douglas-fir (Fd8, >100 yrs). Preharvest through 4/5 season postharvest conditions. Pheromone traps were baited with a frontalin, MCOL, seudenol lure specific to Douglas-fir beetle

Preharvest Post 1 Post 2 Post 3 Posl 4/5 (oc = 0.05). 1.7a) Species # (S), b) Harvest Year Margalef's d, c) Peilou's J', d) Brillouin, e) Shannon's H', f) 1-Simpsons g) Fisher's a, h) Taxonomic diversity 8, i) Taxonomic distinctness 8*.

102 intermediate levels in 3ra (98.80) and 4/5tn years (88.20). The trend observed in control data was similar to that of pheromone-baited data. Single replicate data points for 1st and 2nd season postharvest conditions did not fall within the confidence intervals of preharvest control data, and the post 1 data point was an outlier to all other control treatment years. Two-sample T-test of difference

= 0 (vs not = 0) resulted in control data being not significantly different from baited for comparable preharvest T-value = 1.12, P-value = 0.299; post 3 T- value 0.85, P-value = 0.426; and post 4/5 T-value = .069, P-value = 0.523) treatment years.

Margalef's (d).

Data for Margalef's index (d) (Figure 3.3b, Table 3.8) followed a similar trend to those observed for species number (S) (Figure 3.4a). Mean index values for baited and control data were lowest in preharvest sites (7.02, and 10.38 respectively). In pheromone-baited sites diversity increased to a peak of

15.03 in 2nd year postharvest, followed by a decrease to intermediate levels in

3rd and 4/5th years (12.37 and 13.39 respectively). The trend observed in baited data was similar and not significantly different to that of control data for comparable preharvest (T-value = -2.56, P-value = 0.037), post 3 (T-value -

1.90, P-value = 0.099) and post 4/5 (T-value = -1.66, P-value = 0.140) treatment years. Single replicate data points for post 1 and 2 years did not fall

103 within the confidence intervals of preharvest control data, and the post 1 result was an outlier to all other control treatment years.

Pielou's J'.

Mean index values (J') for baited and control data (Figure 3.3c, Table 3.9) were significantly different for preharvest (T-value = -21.14, P-value = 0.000), post 3 (T-value -23.4, P-value = 0.000) and post 4/5 (T-value = -4.87, P-value

= 0.000) (a = 0.05). Baited data showed low and apparently stable diversity from preharvest through post 2 treatment years (0.13, 0.13, 0.12: pre post 1, post 2 respectively). At post 4/5 treatment years, mean diversity increased to a high of .38. Control data followed a different trend than baited data.

Preharvest sites recorded the highest diversity (0.85), followed by a drop in diversity associated with the single site data point for treatment year post 1

(0.54). The diversity index returned to the preharvest level with the single site data point for treatment year post 2 (0.81), and subsequently stabilized through post 3 and 4/5 data sets. The index value observed for the single post

1 site lay outside the confidence intervals for all other treatments.

Brillouin

Baited and control data for the Brillouin index (Figure 3.3d, Table 3.10) were significantly different for preharvest (T-value = -13.14, P-value = 0.000), post 3

(T-value -17.62, P-value = 0.000) and post 4/5 (T-value = -4.48, P-value =

0.011) treatment years (a = 0.05), and exhibited similar trends to Pielou's J.

104 Baited data showed low and apparently stable diversity from preharvest through post 2 treatment years (0.53, 0.61, 0.60: preharvest, post 1, post 2 respectively). At post 4/5 treatment years, mean baited diversity reached a high of 1.63. In control data, preharvest sites were observed to have moderately high mean diversity (2.95), which dropped to 2.04 with the single site data point for treatment year post 1. The diversity index exceeded the preharvest level with the single site data point for treatment year post 2 (3.53), and subsequently decreased through post 3 and 4/5 data sets. Within control data, no differences were observed between treatment years, though the index value observed for the single post 1, and post 2 sites lay outside the confidence intervals for all other treatment years.

Shannon-Wiener (H'-m)

Two-sample T-test indicated baited and control data for the H'i0 index (Figure

3.3e, Table 3.11) were significantly different for preharvest (T-value = -14.31,

P-value = 0.000), post 3 (T-value -19.92, P-value = 0.000) and post 4/5 (T- value = -4.66, P-value = 0.010) treatment years (a = 0.05). The data exhibited similar trends to Pielou's J', and the Brillouin indices. Baited data showed low and apparently stable diversity from preharvest through post 2 treatment years

(0.23, 0.28, 0.27: preharvest, post 1, post 2 respectively). At postharvest year

4/5, mean diversity increased to a high of 0.075. Preharvest control sites were observed to have moderately high mean diversity (1.46), which dropped to 1.09 with the single site data point for treatment year post 1. The diversity

105 index exceeded the preharvest level with the single site data point for treatment year post 2 (1.67), and subsequently decreased through post 3 and

4/5 data sets. Within control data, the index value observed for the single post

1 site lay outside the confidence intervals for all other treatment years, and the index for the single post 2 site lay outside of the confidence intervals for preharvest and post 4/5 treatment years.

Simpson's (1-A)

Baited and control data for the Simpson's index (Figure 3.3f, Table 3.12) were significantly different for preharvest (T-value = -8.56, P-value = 0.001), post 3

(T-value -19.29, P-value = 0.000) and post 4/5 (T-value = -3.78, P-value =

0.019) treatment years (fx = 0.05), and exhibited similar trends J', H'10, &

Brillouin indices. Baited data showed preharvest mean index values of 0.24 decreased to a low of 0.17 in post 1 data. From post 2 through post 4/5 treatment years mean diversity followed an increasing trend to a high of 0.94.

In control data, preharvest sites were observed to have moderately high mean diversity (0.94), which dropped to .084 with the single site data point for treatment year post 1. The diversity index returned to the preharvest level with the single site data point for treatment year post 2 (0.95), and subsequently stabilized through post 3 and 4/5 data sets. The index value observed for the single post 1 lay outside the confidence intervals for all other treatment years.

106 Fisher (op.

Mean Fisher (rx) (Figure 3.3g, Table 3.13) index values for baited data were significantly lower than control for preharvest (T-value = -4.92, P-value =

0.008), post 3 (T-value -6.07, P-value = 0.001) and post 4/5 (T-value = -3.28,

P-value = 0.014) treatment years (fx = 0.05). Baited trap data showed the lowest mean index value in preharvest conditions (9.50), and followed an increasing trend through successive postharvest treatment years to a high of

25.01 at post 4/5. Mean control index values were lowest in preharvest sites

(29.28). Single site values increased for post 1 and post 2 treatment years

(36.04 and 46.60 respectively), with mean site values decreasing through subsequent years to an intermediate mean index value of 41.03 for the post

4/5 treatment year. Within control data no differences were observed between treatment years, though index values observed for post 1 and post 2 sites lay within the confidence intervals for all treatment years.

Taxonomic Diversity (5) .

Baited and control data for the Taxonomic Diversity index (Figure 3.3h, Table

3.14) were significantly different for preharvest (T-value = -12.95, P-value =

0.000), post 3 (T-value -17.73, P-value = 0.000) and post 4/5 (T-value = -3.20,

P-value = 0.033) treatment years (a = 0.05), and exhibited similar trends to J',

H'10, Brillouin, & 1-Simpson's indices. Baited preharvest mean index value of

14.22 decreased to a low of 11.29 for post 1 data. Diversity then followed an increasing trend to a high of 36.04 in post 4/5. In control data, preharvest sites

107 were observed to have moderately high mean diversity (67.04), which dropped to 52.75 with the single site data point for treatment year post 1. The diversity index exceeded the preharvest level with the single site data point for treatment year post 2 (69.71), and subsequently decreased through post 3 and 4/5 data sets. Index values observed for the single post 1 and Post 2 sites lay outside the confidence intervals of all other treatment years.

Taxonomic Distinctness index (5 *).

Index values based on taxonomic distinctness were the most unique and without trend for both baited and control data (Figure 3.3i, Table 3.15). Across treatment years, only post 4/5 data yeilded significant differences in mean index values (T-value = 4.46, P-value = 0.007) between baited and control data (a = 0.05). Baited sites had the lowest mean index value in preharvest conditions (63.01) and the highest mean index value in post 4/5 treatments

(71.49). Mean index values for control data ranged from high of 71.24 for preharvest sites to a low of 66.16 for post 4/5 sites. Single site values from post 1 and 2 treatment years exceeded the confidence intervals observed for baited data, but again the data presented no clear trend.

Species abundance - baited vs. control

Wilcoxon Rank-Sum analysis in preharvest, post 3 and post 4/5 conditions identified a total of 17 instances where species abundance was significantly different between baited and control sites. Reproducibility across treatment

108" . years varied from one to five consecutive treatment years depending on the species. Table 3.16 shows that 12 species made up the 17 instances, with 9 species occurring once, 1 species (Anaspis sp.) occurring twice, and 2 species (Dendroctonus pseudotsugae and Thanasimus undatulus) occurring in all three assessed treatment years.

Table 3.16. Species of flying Coleoptera statistically more or less abundant between pheromone-baited and unbaited control sites in preharvest, 3rd, and 4/5th season postharvest conditions. Forest habitat was mature to overmature interior Douglas-fir undar attack by the Douglas fir beetle prior to harvesting. Pheromone-baited traps were baited with a frontalin, MCOL, seudenol lure for Douglas-fir beetles (Dendroctonus psuedotsugae). Harvest Rank Increased Rank Year Increased abundance Sum abundance Sum Pheromone-baited Value - Critical Unbaited Value - Critical traps Baited Value control traps ControlValue Preharvest Dendroctonus 0 14 Drasterius 11.5 14 pseudotsugae debilis SCOLYTIDAE ELATERIDAE Thanasimus undatulus 1 14 eanthophlax 9.5 14 CLERIDAE irificus ERAMBYCIDAE Anaspis sp. 14 14 STAPHYLINIDAE Polygraphus rufipennis 13.5 14 SCOLYTIDAE

Rhizophagus dimidiatus 13 14 RHIZOPHAGIDAE Rhinosimus viridiaeneus 13 14 CURCULIONIDAE ^season Dendroctonus 0 8 Hypnoidus 7 8 postharvest pseudotsugae bicolor SCOLYTIDAE ELATERIDAE Thanasimus undatulus 0 8 CLERIDAE Calitys scabra 5 8 (TROGOSTIIDAE)

Anaspis sp 7.5 8 (STAPHYLINIDAE) ~4/5m Dendroctonus 0 3 Hadrobromus 3 3 season pseudotsugae americanus posffrarvesfSCOLYTiDAE CARABIDAE Thanasimus undatulus 0 3 CLERIDAE Trypodendron lineatum 2 3 SCOLYTIDAE

109 Species trends

The final analysis was trends species abundance across treatment years from preharvest sites through 1st, 2nd, 3rd, and 4/5th season postharvest conditions.

Of 586 species identified from pheromone traps, 351 spp followed one of four trends: Increasing mean abundance/site (49 species); decreasing abundance

(42 species); increasing then decreasing (253 species); or decreasing then increasing (7 species) from preharvest through postharvest conditions (Figure

3.4). An additional 103 spp followed no discernable trend. Species with single occurrences (132) were omitted from analysis. Trends observed from control species resulted in similar species distributions. Of 385 species identified, 187 species were found to follow a trend; 54 species increased in mean abundance/site; 17 species decreased; 96 spp increased then decreased in mean abundance/site; while 20 species decreased then increased from preharvest through postharvest conditions (Figure 3.5). An additional 101 spp followed no discernable trend. Species with single occurrences (97) were omitted from analysis.

110 100000

Preharvest Post 1 Post 2 Post 3 Post 4/5 Harvest Year

Figure 3.4. Trends in flying Coleoptera trapped in Douglas-fir beetle pheromone- bajted Lindgren funnel traps under preharvest and postharvest conditions. Trends are presented as the sum of mean abundances/site for species within a trend. 354 spp followed 1 of 4 trends: Increasing mean abundance/site i(49 species); decreasing m (42 species); increasing then decreasing ^ (253 species); or decreasing then increasing >K (7 species) from preharvest through postharvest conditions. 103 spp (not shown) followed no discernable trend. Species with single occurrences (132) were omitted from analysis.

10000

0.1 -I , 1 . 1 1 Preharvest Post 1 Post 2 Post 3 Post 4/5 Harvest Year

Figure 3.5. Observed trends in flying Coleoptera trapped in preharvest and postharvest conditions trapped by unbaited Lindgren funnel traps. Trends are presented as the sum of mean abundances/site for species within a trend. 186 spp followed 1 of 4 trends: Increasing mean abundance/site A(54 species); decreasing a (17 species); increasing then decreasing £ (96 species); or decreasing then increasing JK (20 species) from preharvest through postharvest conditions. An additional 97 spp (not shown) followed no discernable trend. Species with single occurrences (101) were omitted from analysis.

111 All four trends observed in pheromone-baited data were found to occur in unbaited control data in similar proportions, including the number of species not observed to follow a trend (103,97: baited, control), and the number of species with single occurrences (132,101). The largest difference between baited and control data within a trend was observed in the 'increased then decreased' trend, where baited data more than doubled in the number of species holding the trend (253 species compared to 96 species for unbaited data). A list of species assemblages within each trend is presented in

Appendix III.

Discussion

The size of the data set resulting from this study creates a potential for interpretation and discussion at many levels; from species, to families, to succession, to CWD utilization, to the efficiency of biodiversity measures, in the context of both harvesting and pheromone baiting. In an effort to address a range of discussional elements while retaining some focus, study results will be represented from two main perspectives on the flying beetle community associated with harvesting and pheromone baiting in beetle attacked northern interior Douglas-fir:

1) Assessing the effect of harvesting on diversity across treatment years

as measured by unbaited and baited funnel traps.

112 2) Assessing the impact of pheromone baiting on the trap catch of flying

beetles as represented by changes in flying beetle composition and

abundance between baited and control traps within treatment years.

The format of discussion will consist of a brief summary review of results specific to either harvesting or pheromone baiting followed by an interpretation of the results with respect to current research/theories. Single species assessments and family comparisons are integrated into the discussion where possible to illustrate interpretations, though a comprehensive assessment of individual species will not be completed beyond the summary table (Table 3.1) and the presentation of trends (Appendix III).

Impact of Harvesting

The analysis of pheromone-baited and unbaited control data, from preharvest through 4/5th season postharvest habitat conditions, indicates that harvesting affects measurable changes on flying beetle communities associated with beetle attacked interior Douglas-fir habitat. A non-linear increase in species richness and total abundance after harvesting resulted from the combined impact of variable abundance patterns from individual species. A temporary increase or "surge" in abundance after harvesting was the dominant response of flying beetle species, with more than half of the species assessed (in both pheormone baited and control traps) exhibiting their highest abundance within

113 the first two years following harvesting. This trend was found to be one of five identifiable trends:

1) surging abundance (increasing then decreasing), 2) increasing abundance, 3) decreasing abundance, 4) depressed abundance (decreasing then increasing), 5) and no trend

the majority of species could be classified within these trends based on changes in mean abundance/site from preharvest through postharvest conditions. The five abundance trends were present in similar proportions for both pheromone baited and control traps, and the observed changes in species abundance and resulting trends are thought to reflect species responses to changes in habitat suitability.

Surging species abundance

The trend of 'surging abundance' (represented by an increase in species abundance within the first two years following harvesting, followed by decline) is thought to reflect a group of species specifically responding to the disturbance event of harvesting. The abundance patterns of this group follow that of the Douglas-fir beetle, and their lifecycle associates with CWD immediately following harvest disturbance.

114 Invertebrate utilization of CWD was described by Ehnstrom in 1979 (presented to english audiences in Heliovaara and Vaisanen (1984)), and consists of four successional phases (Figure 3.6), the first of which appears to correspond with stage 1 of CWD decay (Maser et al. 1988), as well as the 'disturbance response' guild (increasing then decreasing abundance trend) observed in this study.

Figure 3.6. Succesive phases on the invertebrate community exploiting dead wood. Phase A - a short-term stage mostly with species feeding on bark, such as Scolytidae, Cerambycidae, and species living in their cavities, as well as parasites and predators. Phase B - a rather short stage with species mainly living under bark and in the surface layer of timber, and species associated with fungi. Bark becomes loose and falls off the stem. Phase C - a long term stage which can take several decades, mostly with wood inhabiting species. Phase D - A very long stage during which many wood inhabiting species are replaced by species living under the shelter of decaying logs. Note that much variation occurs depending on the tree species and geographical areas in question (according to Ehnstrom, as presented in Heliovaara and Vaisanen 1984)

115 Ehnstrbm's phase A, CWD's stage 1, and the disturbance response guild observed in this study are comparable in three aspects: CWD condition, temporal occurance, and species composition.

The condition of the decaying wood associated with Phase A and CWD stage

1 is described as the wood having bark still intact on the stem (Heliovaara and

Vaisanen 1984, Maser et al. 1988). Temporally, phase A is shown as having a duration of two years however, Ehnstrom noted that much variation occurs depending on tree species and geographical areas, and Douglas-fir is known to resist decomposition (Carpenter et al. 1988). In this study, stumps and large debris resulting from harvesting were observed to retain bark through the duration postharvest trapping though surveys to assess the condition of smaller stems and rates of decay were not included in study parameters.

According to Ehnstrom's model of CWD, Phase A utilization by insects is defined by species feeding on bark, such as Scolytidae, Cerambycidae, and species living in their cavities as well as predators and parasites. This definition corresponds with the general community descriptions of stage 1

CWD decay described by many researchers (see Caza 1993, Dajoz 2000).

Scolytids, cerambycids and burprestids are present in the study data and are strongly represented in the 'surging abundance' trend described earlier. The trend contains over half of all of the scolytid, cerambycid and buprestid species observed in the study (Appendix III - Table 3). In addition to expected

116 families, relatively large species representations were observed from unexpected families, most notably; Carabidae, Elateridae, and Staphylinidae.

The number of species from each family and their similar distribution among observed trends to that of established scolytid, cerambycid and buprestid families suggests that carabids, elaterids, and staphylinids may have a larger role in postharvest habitats associated with CWD than previously considered.

A survey of some of the more abundant species occuring immediately following harvesting, includes a number of known primary and secondary attacking species including:

the Douglas-fir beetle (D. pseudotsugae) the target species of the study, known to be a primary attacking bark beetle on Douglas-fir (Bright 1976), Hylastes nigrinus - a bark beetle known to feed and complete reproductive cycles in association with the roots of recently dead or dying Douglas-fir (Rudinsky and Zethner-Moller 1967). Asemum striatum - a Cerambycid that reproduces in the sapwood of recently killed Douglas-fir, true firs, larch, spruce and pine (Furniss and Carolin 1977), Trypodendron lineatum - a holearctic scolytid species that attacks the sapwood of any in its range (Bright 1976). Ambrosia beetles are almost exclusively associated with disturbance events. Penetrating fresh downed logs, shaping community development and biological activity within a changing physical and chemical environment (Schowlater ef al. 1988, Byers 1995). Melanophila drummondi - a secondary (and occasionally primary) attacking buprestid beetle most frequently found on Douglas-fir, true firs, spruce, western hemlock and western larch (Furniss and Carolin 1977).

Life-history information above confirms a multi-species assemblage of flying beetles associated with the death and initial decay of a Douglas-fir tree.

Studies of species assemblages associated with pine beetle attack (and the

117 resulting pine mortality) indicate that species presence may not be strictly limited or defined by a habitat association, and some species may be occuring in the context of interspecific associations. A comparison of beetle species from this study and those known to associate with Dendroctonus brevicomis in beetle attacked western pine (Stephen and Dahlsten 1976), and D. ponderosae (monticola Hopkins) in attacked yellow, western white, and logepole pine (De Leon 1934) indicates a number species common to both habitats, including:

Enoclerus sphegeusb-a clerid predator on Dendroctonus spp. Cucujus clavicepsc - Flat bark beetles who's larvae and adults are entomophagous predators on cambium dwelling insects (Smith and Sears 1982). Enicmus tenuicornis - a species of Lathridiidae with an unknown association to pine and Douglas-fir. Xylita laevigatabc and Scotochroa basalisb - melandryid beetles with an unknown association to pine and Douglas-fir. Bius estriatus b - a tenebrionid beetle with an unknown association to pine and Douglas-fir. Spondylus upiformis - a cerambycid with limited lifehistory information that has been observed to bore into the roots of (Furniss and Carolin 1977). Polygraphus rufipennisbc - A scolytid known to be associated with dead and dying spruce, pine and larch (Furniss and Carolin 1977), not known to be associated with Douglas-fir. Gnathotrichus retususbc - A scolytid that colonizes and reproduces in the sapwood of Douglas-fir, true firs, and pines (Furniss and Carolin 1977, Liu and McLean 1993). Hylurgops rugipennis c - A scolytid that breeds in the root crown or stumps of pines, spruce, Douglas-fir, and western hemlock (Bright 1976, Furniss and Carolin 1977). Ips latidensc - breed in the tops or limbs of weekened or dying pines (Furniss and Carolin 1977), Douglas-fir and hemlock (Bright 1976). Orthotomicus caelatusbc - A sapwood colonizing scolytid whos hosts are thought to include all species of conifers within its range (Bright 1976).

118 Abundance records from this study for 10 of the 12 species indicate the species abundance follows the surging abundance trend characterised by the

Douglas-fir beetle for either baited traps (b), control traps (c) or both (bc). In all cases but one {Polygraphus rufipennis), available life history information indicates an association between these species and either the Douglas-fir beetle or Douglas-fir trees. When observed in pine habitat these species are considered associates with resident Dendroctonus species (D. brevicomis or

D. ponderosae (monticola Hopkins)). Given the similarities in habitat utilization between Dendroctonus spp, the interspecific associations defined in pine habitats may also occur in Douglas-fir in association with the Douglas fir beetle. This suggests the presence of a group of disturbance response species that utilize more than one host and may associate with more than one primary attacking beetle species. Representing a generalist species component of a larger, "phase A", disturbance response guild of an unknown size with an unknown extent of species assocations.

Species richness is perhaps the most significant differences between

Ehnstrbm's phases of CWD utilization, the known associates of stage 1 of

CWD decay, and this study. Ehnstrom's model gives no indication of the number of species associated with each phase of succession in CWD. It only indicates that the total number of insects in each phase will be equal at the peak of the phase. Studies of insect assemblages associated with stage 1 of

CWD have found from 30 to 86 species of Coleoptera from 12 to 26 families in

119 association with recently felled spruce (Gara et al. 1995), western pine beetle attacked, dying ponderosa pine (Stephen and Dahlsten 1976) and mountain

pine beetle attacked lodgepole pine, yellow pine and western white pine (De

Leon 1934). These species assemblages are smaller than the 96 species (28 families) classified in the 'disturbance response guild' from unbaited trapping results, somewhat less than the 254 species (49 families) classified in the disturbance response guild from pheromone baited data, and substancially less than the 595 species recorded from traps in 1st season through 4/5th season postharvest conditions. Reasons for the increase in the species number associated with the 'disturbance response guild' and post harvest trap data of this study are uncertain, but may include: 1) a greater number of species associated (both directly and indirectly) with CWD than previously observed; 2) the presence of a naturally higher species complement associated with Douglas-fir CWD in stage 1 of decay compared to previously assessed spruce and pine assemblages; 3) in the case of pheromone baited data - the presence of a non-target pheromone influence on species data; or lastly 4) the presence of flying beetle species associated with other plant species or successional factors not associated with CWD. The potential influence of non-CWD assocations and other successional factors are presented in the remaining species abundance trends.

120 Decreasing species abundance

Losses or declines in temperate forest beetle species following severe habitat alteration is thought to reflect species preferring (or limited to) preharvest habitat conditions (Lavallee 1999, Halme and Niemela 1993). A decreasing trend was observed for 42 species in baited traps and 19 species in unbaited traps (Appendix III - Table 1). A review of species life history information from study data identifies three examples of habitat limited species with declining abundance following harvesting: One species limited by stand conditions, the second limited by prey (changing because of stand conditions), and the third species limited by a combination of habitat and prey.

Quedius criddlei (Staphylinidae) is thought to represent an example of a species limited by habitat association. The life habits of this species are relatively unknown, but national collection material bear habitat labels indicating a strong association between the species and damp decaying habitats (collection tags include "under board on meadow, rotten log

Pseudostuga taxifolia, rotten log Abies grandis,..."), (Smetana 1971). Based on this information the Staphylinid may not be limited to mature, old growth

Douglas-fir, but appears to be associated with advanced decay - a condition that characterizes old growth temperate forests (Vaisanen et al. 1993).

A second species for consideration is a bark beetle predator first observed and discussed in Chapter 2. Under preharvest conditions Rhizophagus dimidiatus

121 (Rhizophagidae) occurred in significantly greater abundance in baited traps

(Table 2.3 and Table 2.6, Chapter 2). The beetle is thought to be predatory on

Scolytids (Deyrup and Gara 1978), and although the species was present in postharvest trapping conditions its abundance pattern did not follow the trap abundance observed for the Douglas-fir beetle. Assuming adequate species distribution, and similar preharvest and postharvest trapping capabilities, the reason for the postharvest decrease in abundance may be that the predator is unable to utilize post harvest beetle conditions to the same extent as it's prey.

This combination of habitat and prey limitation is thought to occur in Cucujus claviceps, which has been described as a generalist predator that preys on scolytids and cerambycids in shaded conifers (Deyrup and Gara 1978).

Decreasing abundance of the predator in pheromone-baited traps supports a combined prey + habitat limitation to species abundance. However, unbaited trapping data generated conflicting results to the expected habitat limitation.

In unbaited traps a temporary increase in postharvest trapping abundance

(Table 3.5) was observed in first and second season postharvest conditions, indicating that the species is present in postharvest conditions. Furthermore, the surge in species abundance observed immediately after harvesting suggests that predator abundance follows prey abundance and the predatory cucujid may be more accurately described by the disturbance response guild.

122 Increasing species abundance

Increasing trends were observed for 51 species in baited traps and 55 species in control traps (Appendix III - Table 2). This trend is consistent with expectations of increasing diversity generally associated with seconday succession (Perry 1994) and should include species associated with plant succession (McLeod 1980), stand structure and microhabitat development

(Southwood et al. 1979), as well as the community associated with intermediate and long term decomposition of CWD (Heliovaara and Vaisanen

1984).

Species associations with open habitats, meadows, or forest edges are known to exist for a number of species observed in the study including; Carabidae

Bradycellus nigrinus (Lindroth 1968) Carabidae, Ctenicera spp. and Ampedus spp (Elateridae). Elaterids are the most dominant family reflecting an increase in species abundance following harvesting (Appendix III - Table 1). They also dominate the most abundant species in 4/5th season postharvest sites (Table

3.6). Habitat associations of elaterids observed in this study are largely unpublished, but some species (Ctenicera aeripennis (syn. Selatosomus aeripennis) and C. resplendens (Wilkenson 1963) are known to be limited to meadow or grasslands, while other species are associated with forested habitat with margins or openings (Ctenicera umbricola, C. pudica, C. propola columbiana, Ampedus occidentalis (Paul Johnson, personal communication,

123 2002)).2 With respect to CWD successional associations, the Cerambycid species (Callidium cicatricosum) is known to attack dead and dying conifers and is noted for it's utilization of air-dried lumber and seasoned stems as long as the bark remains attached (Furniss and Carolin 1977).

Depressed species abundance

The trend of a depressed species reponse characterised by an initial decline followed by an increase in species abundance is the least represented in terms of species number for baited sites (7 species) and slightly more represented in control sites (20 species) (Appendix III - Table 4). A temporary reduction in species abundance immediately following habitat disturbance could be the response of a habitat generalist, though this description is vague at best and assumes that some aspect of the harvesting event caused a temporary reduction in species abundance. Life history information for the species complement associated with this trend is very limited. Species information has been found for only one species - Calopus angustus

(Oedemerinae) - known only to breed in the wood of dead pine, fir, cedar, willow and cherry (Furniss and Carolin 1977). To complicate matters further, many species present in the trend from unbaited data report depressed abundance in 1st and 2nd season post harvest conditions. Conditions respresented by single site replication, increasing the probability that the observed trend in unbaited species, is influenced by sampling variation.

2. Dr. Paul Johnson, Insect Research Collection South Dakota State University 124 Harvesting summary

Individual species responses combine to create a dynamic community response. Four trends are presented in an attempt to describe changes in species abundance and composition, each trend composed of species whose life histories are highly variable in nature. When life history information is applied to the data, a number of species fit within the context of the trends described above. However, some species' life history information does not fit and we are left to wonder what factors are responsible for the observed departure from expectation. Some species fit either approximately, or with one data set (pheromone baited or unbaited) and not the other. The range in species' life histories is a complicating factor in this study, but it is not unexpected. Niche separation has many dimensions and is a fundamental part of multiple species utilizing resources within a habitat (Krebs 1994). By definition, the development of niches will lead to species specific responses such as those observed this and other studies of forest coleoptera (Niemela et al. 1993).

It should be noted that this presentation of species and trends makes no attempt to define the exact complement of species responding to a harvest event, nor is it this author's intention to present expectations, or define the full range or the exact nature of species associations. The lack of life history

125 information for all species makes it impossible to assertain with certainty why species are present in a particular abundance pattern. The goal is simply to go beyond the limitations of diversity indices to describe dynamic changes in species composition through pattern development. Patterns that bear similarity to one proposed in the late 1970's to characterize the biogeochemical dynamics of disturbance and ecosystem development through secondary succession.

Bormann and Likens (1979) proposed that ecosystem development associated with anthropogenic disturbance could be described by 4 phases - each defined by changes in biomass, species composition, and biogeochemical function (Figure 3.7).

Time Figure 3.7. Phases of ecosystem development after clear-cutting of a second growth northern hardwood forest. Phases are delimited by changes in total biomass accumulation (living and dead organic). It is assumed that no exogenous disturbance occurs after clearcutting (from Bormann and Likens 1979).

126 Following disturbance a relatively brief "reorganization phase", was

characterized by dramatic changes in the forest ecosystem including; decreases in primary production, nutrient uptake, transpiration; increases in decomposition, denitrification, biomass, soil moisture, soil temperature; as well as dramatic changes in nutrient stores and the species composition of vegetation. While Bormann and Likens (1979) make no mention of insect diversity associated with the reorganization phase, the time span of the reorganization phase encompasses the species diversity and temporal abundance patterns observed in this study. Additionally, changes in the physical and geochemical components of the reorganization phase lack the directional progression and relative stability characterized by other phases of forest development. In the context of this study and its assemblage of flying beetles, the reorganization phase contains and reflects the sum of all species interactions. These are interactions that when taken as a whole cross structural, compositional and functional levels of biodiversity, and include flying beetle species associated with disturbed habitat, old growth habitat, regeneration habitat, transients, and those unaffected by habitat change.

These dynamics are described as a "composite of variable ecosystem processes" characterised by a complexity that is consistent with multiple insect communities and/or species groups such as those observed in this study. It is interesting to note that using net biomass change as the indicator, the duration of the reorganization phase is estimated at 15 years for temperate hardwood

127 forests (Borman and Likens 1979). This duration extends beyond the time frame of this study to encompass successional phases A, B, and the beginnning of phase C of CWD insect colonisation (Heliovaara and Vaisanen

1984) and the first 2 stages (0-18 years) of coarse woody debris decay established for Douglas-fir (Maser et al. 1988).

Flying beetles associated with Douglas-fir beetle attack, the harvesting of interior Douglas-fir, and the utilization of CWD in forest succession can be assessed as a single community, or a combination of smaller communities.

Either way, the results of this study indicate that the harvesting of beetle attacked trees affects significant changes in the species diversity of flying beetles. This result occurred for both pheromone-baited and unbaited trapping conditions. However, the extent of change varied between baited and unbaited conditions, indicating a difference in the trapping efficacy of pheromone baited and unbaited Lindgren funnel traps.

Pheromone Effect

Results show that baiting with synthetic pheromone lures for Douglas-fir beetles changes species diversity of baited trap catches compared to unbaited trap catches under both preharvest and postharvest conditions (Preharvest,

1st, 2nd, 3rd, & 4/5th season postharvest). Indices assessing richness (H\

Brillouin, Fisher's fx), evenness (J'), dominance (1/D), and taxonomic diversity

(5) produced consistent results of decreased diversity associated with

128 pheromone baited trap assemblages. While decreased diversity is generally held to be synonymous with ecological quality (Magurran 1988), the significance of the results are not based in any assessment of quality. Rather the measures indicate significant changes in species presence and relative abundance resulting from pheromone trapping across a range of habitat conditions. The difference is observed for indices emphasing rare species

(richness, and taxonomic diversity), common species (dominance), or the relative distribution of species within the sample (evenness). In contrast, the range of indices designed to assess species richness did not show uniformity in results.

Species richness (S) and Margalef's indices suggest that Douglas-fir beetle pheromone lures do not change the number of species observed in the data pool. However, closer analysis suggests that this result is likely an artifact of low sampling effort, emerging in the form of a species-area curve from indices highly sensitive to sampling effort. Species richness (S) and Margalef's indices are highly sensitive to sampling intensity while all other indices are characterized as having moderate, low, or no sensitivity to sample size (Clarke and Warwick 1999, Magurran 1988, Pielou 1969).

The measure of taxonomic distinctness (5*) is unique and the results generated by the index are in stark contrast to all other diversity indices. The index results in a mix of trends with errors so large, that baited and unbaited

129 trap catches are statistically indistingushable in all treament years except post

4/5 (with baited traps exhibiting greater diversity than control traps).

Taxonomic distinctness (5*) is a univariate index that attempts to capture phylogenetic diversity, or the taxonomic relatedness of species occuring within a sample by calculating the average 'taxonomic distance' between all pairs of species within a community sample (Clarke and Warwick 1998, 1999).

Reduced taxonomic distinctness has been associated with increased stress and decreased trophic diversity (Warwick and Clarke 1995, 1998). In this study, 5* would suggest that there is no difference in taxonomic distinctness between pheromone-baited and unbaited trapping, or between preharvest and post harvest insect communities. The reason for the observed lack of distinctness within trapping years may be owing to the presence of a single species pool with a high proportion of common and abundant species within baited and unbaited data. Interpreting the observed differences in distinctness between trapping years is less clear because the index does differentiate between either 'end' of the assessed harvest conditions (preharvest, 1st season postharvest conditions and later 3rd and 4/5th season postharvest conditions), but is unable to differentiate between adjacent harvest years.

Under conditions of epidemic beetle attack and the onset of a beetle associated disturbance event, it may be that the index is dominated by small number of species (Douglas-fir beetles and associates) that persist as long as habitat conditions allow (in this case from preharvest through second season

130 post harvest conditions), and only reduce their dominance on the index as habitat conditions become unfavorable.

Returning to the significant results observed at the community and species level, pheromone lures for the Douglas-fir beetle tested in this study differentially trap non-target flying beetles in preharvest through 4/5th postharvest conditions. This result, coupled with the disproportionate increase in the number of species exhibiting the 'disturbance response' trend in baited traps following harvesting (253 species in baited sites vs 96 species control sites (Figures 3.4 and 3.5)) suggests that pheromone baiting not only changes trap catch diversity but also magnifies the number of species following the

'disturbance response' pattern of abundance. Interpretation of the results is complicated by a lack of comparable studies on such a large species assemblage, over the range of habitats (harvest conditions) assessed, using this particular pheromone lure, in conjunction with diversity assessment.

Large-scale multiple species, or mutliple family studies of forest insects have interpreted species associations in one of two contexts - either in response to the disturbance of a specific host species (Werner and Holsten 1984,

Hammond 1997), or in response to attack by a specific bark beetle species

(De Leon 1934, Stephen and Dahlsten 1976, Deyrup and Gara 1978, Gara et al. 1995). While both interpretations are appropriate in context, neither perspective addresses the role of semiochemicals in the development of these

131 communities. In southern pine, chemically mediated behavioural interactions have been identified in insect colonization sequences, resource partitioning and predation strategies (Birch et al. 1980; Billings and Cameron 1984;

Billings 1985, Kohnle and Vite 1984). In Douglas-fir, a recent study by Peck et al. (1997) assessed multiple species responses of scolytid beetles to a pheromone lure similar (though not directly comparable), to that used in this study. Lures trapped over 44 species of scolytids, and many species were thought to be attracted to one or more semiochemical components. With the lack of research, interpreting the impact of pheromone lures on community structure is necessarily theoretical in nature, though it is based in forest ecology, and on the role of semiochemicals (both singly or as a part of a multiple chemical concert) in the development of the forest environment.

Bark beetle pheromones are volatiles resulting from metabolic processing of beetle steroids and the detoxification of terpenes originating from the host tree

(White et al. 1980). Cross-feeding experiments in Dendroctonus spp. indicate conserved metabolic pathways occur between species within the genus

(Libbey et al. 1985), suggesting that host tree biochemistry is a major determinant of pheromone composition for primary attacking species.

Following beetle attack, host condition (and it's associated biochemistry) is reflected in bark beetle pheromone production, resulting in distiguishable combinations of specific and parsimonious pheromones with varying influence on beetle activity (Libbey et al. 1985). Variation in semiochemicals including

132 host-derived kairomones, aggregation pheromones, nonhost volatiles and gustatory cues create a dynamic sensory map of the forest environment for any species that can read and interpret the signals (Huber et al. 2000). The development of synergistic effects utilizing multiple pheromones and kairomones increase messages of habitat availability, and are thought to be part of an adaptive strategy for beetle species utilizing ephemeral resources

(Hedden et al. 1976). The ability of Douglas-fir beetle pheromone lures to alter the diversity of flying insects in both preharvest and postharvest conditions may be related the role of insects, particularly bark beetles of the genus Dendroctonus, in the production of ephemeral resources through temperate forest disturbance events.

In ecology it is a widely held tenet that animal species richness is tied closely to plant successional patterns (Perry 1994), and the faunal diversity of insects reflects habitat conditions (Lattin 1993). The relationship between the plants and insects is integrated to the extent that the sequence of establishment, proliferation and decline of plant species will determine the sequence of establishment, proliferation, and decline of host-specific insect species. The extent of interaction is such that changes in insect assemblages, induced by vegetation change, appear to regulate fundamental ecosystem processes such as decomposition, energy flow, and nutrient cycling (Schowalter 1981 and 1985, Wood 1982, Edmonds and Eglitis 1989) - processes that ultimately infuence the rate and direction of succession. Because of their capacity to kill

133 living trees over large areas, bark beetles (including the Douglas-fir beetle) influence the age, size, and species distributions of forest flora, and thus are a significant factor in forest succession (Wood 1982, Huffaker et al. 1984).

When bark beetles tend to specialize on a single dominant host, their ability to create and respond to disturbance events strongly controls both the rate and direction of succession (Haack and Byler 1993). These disturbance processes generate and maintain gap dynamics that impact biodiversity, wildlife habitat, scenic quality, recreation, timber volume, and other forest resources (Lundquist 1995). In the absence of fire, the Douglas-fir beetle has the potential to determine the direction of secondary succession, and in doing so, can affect the development of the species complement (both plant and insect alike) associated with disturbance/succession events.

The role of the Douglas-fir beetle in locating, utilizing and even creating disturbed habitat through semiochemical production and perception supports the potential for non-target disturbance associated beetle species to utilize

Douglas-fir beetle pheromones as kairomones. The results of this study indicate that non-target species are disproportionately trapped by pheromone lures, although the full extent or nature of species associations are unknown, and cannot be determined through the results of this study. Species abundance assessments within treatment years identified 12 species and 17 instances of significant trapping bias towards either baited or unbaited traps.

For the non-target species differentially occurring in baited or unbaited traps,

134 the occurrence of species with significantly different abundance across treatment years varied from one to five years.

Of the 12 species listed, aggregation responses have been established for the

Douglas-fir beetle and its predatory clerid beetle, Thanasimus undatulus (Ross and Daterman 1995). The remaining species have never been tested for a response to Douglas-fir beetle pheromone components, and for most of those species there is not enough life history information available to determine the exact cause of disproportionate trapping. A number of potential associations are present and include direct and indirect associations based on non-target species utilizing Douglas-fir beetle pheromones as kairomones to secure resources (see Chapter 2 discussion). As mentioned, the most notable kairomonal response to Douglas-fir beetle pheromones was observed in the clerid, Thanasimus undatulus. The species was observed in significantly greater abundance in baited traps across all preharvest and post harvest conditions. This consistency of result was not observed in the other 10 non- target species that showed significant abundance. The reason for the variability in species occurrence and significant abundance may be the result of a number of factors including: natural species distributions, variable physiological and behavioral responses to semiochemicals, differential trap influence between preharvest and postharvest conditions and/or variation within the semiochemicals of the lure.

135 Natural variation

The spatial distribution of insects in the forest is non-uniform and is determined by the behavior and life history requirements of each insect relative to its environment (Dajoz 2000). Single species assessments have attempted to accommodate for any lack of a normal distribution with a distribution free test (the Wilcoxon Rank-Sum test) (Milton 1992), but the relatively low sample size used in this study (from 5 to 11 replicates for each treatment and habitat (Table 3.5)), may not accommodate variations in natural distribution for all species.

The responses of different species of Coleoptera to semiochemicals in the environment result from a combination of perception and response capabilities. Species of forest beetles are thought to succesfully navigate variable habitat conditions characterised by a high degree of semiochemical parsimony (Huber et al. 2000). Variation in behavioral responses between species is thought to result from semiochemical composition, geometric configuration, enantiomeric specificity, volatile concentration, and the receiver's perceptive ability (Werner et al. 1981, Raffa and Klepzig 1989,

Gries 1992, Ross and Daterman 1998, and Huber et al. 2000, respectively).

The presence of parsimonious chemicals may allow some species to identify and utilize multiple hosts, enforce parapatric distributions (Lanier and Wood

1975), or lead to cross attraction and the misinterpretation of habitat availability such as that described earlier in this discussion for Polygraphus

136 rufipennis. The misinterpretation of semiochemicals from pheromone lures causes disproportionate trapping of non-target, non-associated species in pheromone traps. As with other proposed associations between species and lures resulting from this study, the extent of such 'accidental aggregation' is unknown, though its occurrence must be considered to understand the full potential of semiochemical influence. While the presence of the effect does not detract from the lure's impact on non-target species, it does limit our ability to interpret the total impact of the lure as a reflection of species closely associated with Douglas-fir, Douglas-fir beetles, or their natural pheromone system.

Trapping efficacy

The ability of traps to disperse pheromones and sample beetle populations was discussed in Chapter 2 in the context of standing attacked, old-growth habitat. Under preharvest conditions species were thought to be trapped passively through random flight interception, and actively in response to the visual profile of the trap, as well as in response to the variable pheromone plume. The addition of postharvest data to the sampling structure adds another set of factors to this sampling variation. The potential for passive sampling for flying beetles by the traps is not thought to be affected by a change in harvest conditions as trap size (surface area) and position remain consistent regardless of the habitat conditions. However, active sampling in response to the visual profile of the trap does change relative to the adjacent

137 visual profile of the forest following harvesting, and the removal of the forest canopy changes the development of the pheromone plume.

Under preharvest conditions the black vertical silhouette of a multiple funnel trap appears as one of many vertically oriented landing sites (trees) in the forest immediate surrounding the trap. To any visually orienting species with a preference for a vertical landing site, the funnel trap is just one more place to land. Remove the surrounding forest cover through clear cutting and the pheromone trap is the only dark, vertical profile remaining in the area, giving any visually orienting species with a vertical landing preference just one place to land. This change in profile suggests that the active sampling of species based on visual cues from the trap will increase from preharvest to post harvest conditions, though the extent of the impact is unknown.

With regard to pheromone dipersal, postharvest conditions affect many changes on the development of the pheromone plume. Removal of the forest canopy changes ambient temperature at the height of pheromone release through decreased insulation, and the loss of inversion effects from canopy closure, as well as increased solar radiation (Elkinton and Carde 1984). The impact on pheromone dispersal is that the lure is subject to a greater fluctuation in temperature-mediated diffusion rates. In addition to temperature changes, harvesting even small patches of forest canopy results in altered and increased wind patterns within the harvest area (Perry 1994). Increased

138 winds may then alter the size and shape of the effective sampling area of the

pheromone plume (Turchin and Odendaal 1996), as well as the nature of species responses (Salom and McLean 1991).

The nature of pheromone dispersal is fluid, variable, and impossible to predict, even when habitat conditions remain largely stable. The addition of more complicating variables following harvesting seems overwhelming until you consider the nature of the species under study. The Douglas-fir beetle is a disturbance response specialist. It and other species utilizing disturbed habitats have been interpreting, compensating for, and adapting to inherent variation for thousands of years. While it is reasonable to assert that variable conditions will produce variable results, the extent of that variation is unknown.

Assuming the composition and release rates of pheromone lures are consistent with naturally produced pheromones, the measured species responses should reflect natural conditions.

Systematic Bias

Another potential source of variation in the data can be found in sampling design, and the presence of systematic bias in spatial, temporal, and biogeoclimatic conditions. The uneven distribution of sampling sites for baited and control data across study years and biogeoclimatic subzones (see Table

3.1) creates the potential for biased sampling (the nature of which is discussed further in Chapter 4) - however, the presence abundant species across

139 treatments, habitats and sampling years, in combination with highly significant results, suggests that the potential impact of such sampling bias was not large enough to overwhelm the study results.

Semiochemical variation

As noted in the the methodology, the presence of seudenol in the pheromone lure was not a controlled factor. In field conditions, seudenol was observed to be spontaneously produced from MCOL in the presence of acidic water

(present in release devices from precipitation or condensation of atmospheric water), stabilizing at an equilibrium ratio of 40:60, MCOL:seudenol respectively

(see discussion on Semiochemical composition pg 2.40). The presence of this third semiochemical may have resulted in variable species responses.

However, despite the semiochemical variation (or perhaps because of it) the lure placed in the Lindgren funnel traps displayed a seasonal efficacy for trapping the target species, against which non-target responses were assessed.

Summary

Results of this study indicate that pheromone lures designed for Douglas-fir beetle aggregation can effect changes in the the trap catches of flying beetle diversity in old-growth (endogenous) disturbance conditions, and postharvest

(exogenus) disturbance conditions - up to 3rd and 4/5th seasons after a harvesting disturbance event. Changes in diversity and species abundance result from the disproportionate trapping of an unknown number of non-target

140 species thought to be responding to Douglas-fir beetle pheromone lures as an indicator of suitable habitat or resources. Significant changes in the diversity of flying beetles from both preharvest and postharvest trapping conditions suggest a community response to Douglas-fir beetle pheromones occurs within the context of, and in addition to, a dynamic and changing species assemblage responding to the effects of harvesting.

The total community assemblage gathered by funnel trapping flying beetles from preharvest and postharvest conditions is thought to be the sum of nested groups of species, identifiable by abundance trends (present in pheromone baited and unbaited traps) including: declining old growth specialists, an increasing secondary succession species group; a surging disturbance response species group, transient species group, and an unaffected species group.

The degree and extent of interactions between flying beetles, habitat change, and semiochemicals are complex and dynamic by nature. Although the results of this study do not define the nature or extent of species interactions, they do add information to the search for mechanisms governing community organization and the disturbance response of flying beetle communities associated with northern interior Douglas-fir and the Douglas-fir beetle.

141 CHAPTER IV

Pheromones and Integrated Pest Management

Introduction

Ecosystem management has been defined as keeping forest ecosystems functioning well over long periods of time in order to provide resilience to short-term stress and adaptation to long-term change (Hack and Byler 1993).

To the forest manager working towards a sustainable forest, the goal is to maintain compostion, structure and function (native ecosystem integrity) of forest habitat over the long term (Barnes et al. 1998) - a goal that includes the maintenance of species, communities, and their associated ecosystem processes in the context of harvesting and other resource priorities.

Arthropods comprise almost 85% of the diversity found in a temperate old growth coniferous forest (Asquith etal. 1990), and, with a unique pattern of life far removed from the vertebrate condition, the range of relationships between insects and their environment should be subject to special consideration from an ecological standpoint (Chapman 1955). Insects are considered to be intrinsically linked to ecological functions in forest ecosystems (Miller 1993) - critical contributers to forest diversity, soil fertility, long term forest health and sustainability (Haack and Byler 1993). While insects are an integral part of the forest environment, the impact of their presence is not always in harmony with human needs. In recent years

142 monitoring and managing forest pests has become a major concern for forestry in British Columbia.

Managing insect pests in the forests of British Columbia is a co-operative effort between Forest Health managers and entomologists at the provincial, regional, and district levels. The range of management options for

Dendroctonus beetles includes prevention, maintenance, suppression, and even abandonment of natural beetle populations, depending on available resources and management goals (Anonymous 1995, Shore etal. 1996).

When populations are present and abandoning them is not an option, the main tactics for the maintenance or suppression of beetle populations include

(Shore etal. 1996):

1) Harvesting a. Single tree b. Selective c. Patch cut d. Clear cut

2) Felling and burning of infested trees

3) MSMA of infested trees

4) Debarking of infested trees

5) Pheromone applications in conjunction with options 1-4. a. Monitoring b. Mass-trapping c. Concentration & containment d. Post-treatment mop up e. Attack disruption

143 Pheromone applications that trigger aggregation responses are used operationally in North America for monitoring, mass-trapping, concentration and containment, and post-treatment mop up of infestations (Borden 1994).

Aggregation of the Douglas-fir beetle using a ternary pheromone blend

(containing MCOL, seudenol, and frontalin), at various release rates, alone and in combination with ethanol, has been reported in central British Columbia and the northwest United States (Guthrie and Wieser 1997, Ross and

Daterman 1998, respectively). Under preharvest conditions pheromone lures in traps or tree baits predictably manipulate beetle colonization patterns, allowing for the removal of large numbers of beetles by harvesting attacked trees surrounding the lures (Thier and Patterson 1997, Shore et al. 1990).

Lures released from Lindgren funnel traps can be used to monitor the relative size of bark beetle and predatory beetle populations (Aukema et al. 2000).

Multiple applications of pheromone lures are a promising tool for IPM programs, in part owing to their potential for a "species-specific" effect (Vite and Baader 1990). The results of this study indicate Douglas-fir beetle pheromone lures in Lindgren funnel traps result in the trapping of both target and non-target flying beetle species.

Pheromone lures composed of the semiochemicals (frontalin, MCOL and seudenol) designed to be specific to the Douglas-fir beetle, were observed to disproportionately trap a multispecies assemblage of beetles, changing both the abundance and distribution of species in baited traps over unbaited traps.

144 The observed differences in diversity resulted from, but were not limited to, identifiable species with significantly different abundances between baited and unbaited traps. Pheromone-mediated differences in species diversity were thought to occur in addition to a large species complement resulting from random flight intercept trapping, and physically (visually) mediated trapping conditions. An attempt at life-history reconciliation found most species to have no investigated pheromone history, and limited life-history information available. Of those significantly trapped species with available life- history/semiochemical information, the target species (D. pseudotsugae), one predatory species (7. undatulus) and one habitat-associated species (7. lineatum) were known to be aggregated by one or more of the pheromone component(s)/lure used in this study. One species (R. dimidiatus) is thought to be associated directly or indirectly with dead or dying Douglas-fir, and one species (P. rufiipennis) is not known to be associated with the beetle nor

Douglas-fir habitat (see Table 3.16 and Chapter 3 discussion). The resulting complexity of evaluating the study's significant results creates a line of interpretation without absolute resolution - one that ultimately must address the role of pheromones developing insect communities in the forest environment and the potential for this pheromone lure to measure and manage those communities.

145 Pheromones, Insect Ecology, and Community Development

Despite a wealth of information on the impact of pheromones on a number of primary and secondary attacking scolytid beetles, little is known about the role of the semiochemicals in the development of forest insect communities. It is known that non-target beetle species will use pheromones as kairomones in the search for suitable resources (Allison et al. 2001, Setter and Borden

1992), and that predators of bark beetles will use prey pheromones and other kairomones as part of generalist or specialist strategies to secure not only prey, but locate mates and suitable breeding sites (Kohnle and Vite 1984).

Beyond a limited number of specific studies, the potential for a multispecies effect that would explain the results of this study is largely theoretical but worth reiterating because the potential presence of a multi-species semiochemical influence reflects forest ecology, and as such, impacts management.

The action of semiochemical systems between beetle species within a genus and sub-populations within a species are considered to be highly evolved

(Wright 1958, Bakke and Kvamme 1981, Payne et al. 1984, Lindgren 1992, and Raffa 2001)), and capable of mediating host selection, mate attraction, resource competition, and predation (Hedden etal. 1976; Chapman 1963;

Lanier 1970; Gast et al. 1993; Lessard and Schmidt 1990; Smith etal.

1990; Rankin and Borden 1991: Hermsera/. 1991). Pheromone efficacy appears to be highly fixed in some species (Borden er al. 1980), variable and

146 labile for others (Lanier and Wood 1975, Hermes et al. 1991), adapted by each species to promote reproductive success despite high degree of semiochemical parsimony (Huber et al. 2000). The presence of a multispecies non-target beetle response to pheromone lures for the Douglas- fir could result from any number of potential associations. However, it is considered that the most probable associations would be related to the formation of ephemeral disturbance habitat resulting from the activity of the

Douglas-fir beetle.

As a species associated with succession and the formation of long-term landscape patterns in Douglas-fir, the Douglas-fir beetle plays an integral role in the natural maintenance and development of North American Douglas-fir forests. Forests of Douglas-fir and its' associated insects have been co- evolving in British Columbia since North America's last ice age (Byers 1995).

Assuming some degree of stability in successional processes, the forest community associated with Douglas-fir beetles, disturbance, and gap succession is the product of at least 10,000 years of co-evolution. Coevolved systems are widespread in insects, frequently involving unrecognized relationships (Bush and Hoy 1984), and far more complex than has been considered (Birch etal. 1980).

Both semiochemical theory and the established ecology of the Douglas-fir beetle support the presence of multispecies response to Douglas-fir beetle

147 pheromone components. While neither theory, nor this study is able to determine the extent of the pheromone response, the presence of a non- target response is relevant to the use of pheromone-baited traps in the management of Douglas-fir beetle populations.

Management Impact

In pheromone-facilitated management of Douglas-fir beetles, baited funnel traps are most easily applied to trapping for containment/mop-up, and monitoring, all of which are relevant in the context of a multispecies, non- target influence by pheromone components or lures.

Containment & mop-up

Containment tactics entail the use of pheromone lures in tree baits or traps, under preharvest conditions, to aggregate Douglas-fir beetles in predictable areas (Ross and Daterman 1997, 1998). 'Mop-up' uses lures in tree baits or traps in or near recently harvested areas to aggregate residual pest populations for subsequent trapping or small-scale harvesting. At the site level the effect of containment on beetle species and other organisms is thus two-fold; aggregation of species into an area followed by habitat and species removal. The benefit of a pheromone-harvesting containment tactics is the ability to manipulate the location of bark beetles, and induce beetle attack on trees adjacent to pheromone lures (Ross and Daterman 1997, Prenzel et al.

1999) to maximize the number of beetles removed from the forest for each

148 tree harvested. When applied consistently, the tactic can reduce the amount of harvesting required to manage pest populations (Borden 1994), resulting in intense management in some areas coupled with complete habitat retention elsewhere. Such a management option can then be balanced against resource needs, non-resource objectives, and the ecological impact of the management process.

Managing for forest pest species is not a simple task. The process has been known to diminish pest-caused structural diversity, decrease functional diversity associated with interacting diseases, insect and other disturbance agents, and alter the abundance and distribution of decomposing dead wood

(Lundquist 1995). In the context of a multiple-species response to pheromone lures, conceptual models and empirical data all suggest that the disruption of multispecies interactions will result in an altered ecosystem with positive or negative impacts depending on the ability to fulfill management objectives (Miller 1993). Non-target species responses to Douglas-fir beetle pheromone lures observed in this study are known to include predators

(Thanasimus undatulus), habitat associates (Trypodendron lineatum), and even species not known to be associated with Douglas-fir trees or the

Douglas-fir beetle (Polygraphus rufipennis) (Table 3.16), suggesting a management impact that extends beyond manipulating a single pest species.

However, neither scientists nor forest managers know the full extent of these associations. In addition, the full life histories, ecological roles, natural

149 distributions, or population levels of the vast majority of the species occurring in this study have yet to be described, leaving management implications subject to theory and conservative conjecture. The lack of fundemental information is perhaps best reflected in the confirmation of at least 4 previously unknown species of beetles from this study from an estimated

15,000 - 35,000 undiscovered / undescribed species of insects thought to currently reside in British Columbia (Scudder 1996).

Despite what we don't know, scientists do recognize the need for minimizing non-target interactions. The negative effects of pheromones on natural enemies of the target organism must be minimized if mass trapping is to be used as part of a wide-scale integrated pest management system (Ross and

Daterman 1995). However, these same researchers complicate the impact of pheromone use by advocating simultaneous applications of aggregation and antiaggregation pheromones within the landscape (Ross and Daterman 1994,

Ross 1997), the combination of which is known to alter colonization behavior of Douglas-fir beetles (Hedden and Pitman 1978). Investigations have also been recently published on the potential for manipulating more than one target species at a time (Greenwood and Borden 1999). This would further confound the issue of the impact on associated nontarget species by increasing the number and range of applications for pheromone lures, and in doing so, increasing the potential scale of impact.

150 The issue of scale is critical to understanding the impact of alteration on a community or ecosystem, with short-term, small-scale impacts posing little threat to sustainability (Toman and Ashton 1996). Assessing changes in faunal assemblages, whether influenced by pheromone applications, containment tactics, or other management efforts must consider the scale of disturbance relative to the replacement time of the habitat through the course of succession (McLeod 1980). Stand level management should also consider the redundancy of spatial and temporal patterns to facilitate the dynamics of species succession at all levels (Toman and Ashton 1996). The results of this study identify short-term impacts on species diversity associated with both pheromone trapping and harvesting, but the scale of species manipulation relative to total species abundance in the district, and the development/distribution of future habitat is unknown, and could benefit greatly from further research into the determination of trapping efficacy and species level assessments of effective sampling area (Byers 1993, Byers et al. 1989, Schlyter 1992, Turchin and Odendaal 1996).

Monitoring

Perhaps the most interesting results of trapping with pheromone-baited and unbaited traps in this study was the ability to sample species diversity of a large, multi-species assemblage through both active and passive means.

151 The impact of baited and unbaited trapping from preharvest through post harvest conditions resulted in the sampling of 512 identified species and 129 recognizable species groups (RTU's) from 67 families. The trapped species assemblage identified significant changes in species compostion and abundance associated with disturbance and the onset of secondary succession, and contained within that flying beetle species assemblage was a differential trapping effect by pheromone-baited traps. The net result was a sampling of flying beetles species that crossed structural, functional, and compositional levels of biodiversity, and yet occurred within a relatively well defined set of habitat conditions.

Pheromone bias as context

Patterns in species composition and the relative abundance of species describe diversity at the community level (May 1976). The presence of a community response to Douglas-fir beetle pheromones observed in this study reflects the sum of individual responses of a half million individuals from 641 flying beetle species/RTUs, across a changing ecosystem. Understanding any functional linkages, real or potential, of this species complement requires a specificity of context (Niemela et al. 1992, Jonsson and Jonsell 1999) that can be provided by the Douglas-fir beetle pheromone lure. Pheromone sampling is clearly a biased approach to looking at species assemblages, but in its unorthodox approach the sampling method creates a well-defined context for assessing species diversity. From the context of pheromones and

152 disturbance, functional associations can be proposed and investigated. In this study the context is the communication system of the Douglas-fir beetle, and the forest disturbance that immediately preceeds or follows its presence.

The assessment of biodiversity in an ecological window as narrow as that associated with Douglas-fir beetle attack comes with a defined pheromone lure (chemical composition, release rates) and defined habitat for both preharvest and postharvest conditions. Regardless of a predisposing agent, the development of a Douglas-fir beetle outbreak in mature/overmature forest has been shown to be correlated with specific tree charactersitics including diameter classes, percent host type, stand density, and basal area (Negron et al. 1999; Negron 1998) as well as tree height, phloem thickness, a standardized growth rate to diameter ratio (Shore et al. 1999). Following harvesting, the assessment of beetle habitat is best characterized by two factors: the stand age/species composition and diameter class at which the forest was harvested, and the extent/condition of the resulting CWD. These habitat characteristics include spatial, and temporal, and host condition determinants of species composition (Jakus 1998), as well as presenting a natural complement of host volatiles known to influence beetle species composition associated with kairomonal resposes (Billings 1985). The condition of beetle attack and harvesting can be assumed to produce a complete pheromone bouquet associated with natural conditions of beetle attack and dead/dying trees. This natural bouquet creates an aggregation

153 condition that is likely far more complex than the impact of simplified synthetic

lures, resulting in a local species complement associated with both synthetic pheromone components/lures and natural conditions. Instead of creating a situation of competition between the synthetic lure and natural conditions, the lure and trap benefit from being in the immediate proximity of the natural species complement associated with Douglas-fir beetle aggregation conditions and a developmental stage specific to mature/overmature Douglas-fir stands.

This type of directed or biased sampling system, when placed in appropriate ecological context has the potential to monitor the condition of highly managed ecosystems and associated elements of biodiversity (Noss 1990). An assessment of the relative abundance of target and non-target pheromone mediated species can be made while gathering fundamental ecological information on flight periods, and the distributional ranges of target, non- target, pheromone mediated, and randomly trapped species (see Peck et al.

1997).

The results of this study suggest the potential for utilizing pheromone monitoring in bark beetle disturbance conditions as a method of sampling both target and non-target beetle populations associated with disturbance events.

The potential for monitoring species is appealing from an ecological perspective, but for results to be applicable at a management level, a greater understanding of the impact of pheromone lures on non-target species is required, including a species level understanding of inter and intra specific

154 behavioral variation, regional influences, along with the elucidation of guild associations, habitat associations, and the influence of trapping methods.

Sources of Variation and Experimental Limitations

The results of this study indicate that pheromone lures specific to the Douglas- fir beetle alter the flying beetle diversity of trap catches across a range of habitat conditions. While this result may afford a greater consideration to non- target species in IPM programs, further research is needed to understand the potential sources of natural and experimental varations and address the limitations of this study. If a community of non-target flying beetles does respond to Douglas-fir beetle pheromones the extent of influence will be dynamic, species specific, and subject to many variables. The study identified a total of 12 species with significantly greater abundance in baited sites over control sites, but 9 of these species showed this result in only 1 out of 3 assessed harvest conditions/years. This leaves us to consider whether the occurrence was the result of semiochemical influence as measured, or result of other factors such as inter/intra specific variation to habitat or pheromone conditions, or perhaps variable temporal or spatial distribution.

Species variation

Each species and individual caught in a funnel trap is characterized by a vast range of species specific variation, including natural variations in abundance and distribution, as well as innate behavioral responses to habitat conditions.

155 The spatial distribution of insects in the forest is non-uniform and is determined by the behavior and life history requirements of each insect relative to its environment (Dajoz 2000). Single species assessments have attempted to accommodate for any lack of a normal distribution with a distribution free test (the Wilcoxon Rank-Sum test) (Milton 1992), but the relatively low sample size used in this study (from 5 to 11 replicates for each treatment and habitat (Table 3.5)), may not accommodate variations in natural distribution for all species.

Research has found that species specific behavior is mediated by a large number of internal and external stimuli (Harris and Foster 1995), and pheromone perception can affect variable behavioral responses between species (Payne 1974, Wood 1982) and regional populations (Rudinsky

1966a). Consistency in the geometric production and behavioral responses of the target species from different regional populations to this MCOL, seudenol, frontalin lure create the potential for use of the lure across geographically separated populations. Regional use of a consistent pheromone blend and trapping methods would not only allow for the comparison of separated target populations, but also provide context for assessing the regional variation in non-target species.

Flight behaviors are are also affected, or determined by the physiological condition of responding insects (Atkins 1975, Salom and McLean 1991). The

156 impact of variable physiological condition can be reduced when the temporal duration of sampling encompases a large range of physiological states, such as a single flight season used to delimit sampling in this study. However any reduction in sampling duration (desirable for efficient and cost effective monitoring) will increase the potential for physiological variation.

Habitat variation

Invertebrates are more sensitive to habitat changes in part because they operate at a smaller spatial and temporal scale than vertebrates (Niemela et al. 1993). Pheromone-baited and unbaited control trap catches, from preharvest through 4/5th season postharvest habitat conditions, indicate that harvesting affects measurable changes on flying beetle communities associated with beetle attacked interior Douglas-fir habitat. These observed changes in diversity result from the combined impact of variable abundance patterns from individual species, as they respond to changing plant species composition, temperatures, insolation, visual cues, wind patterns, affecting both the insects and the efficacy of baited and unbaited traps (Elkinton and

Carde 1984, Perry 1994, Turchin and Odendaal 1996).

Identifying the presence and extent of responses for individual species to pheromone baiting and harvesting represents a lifetime of future research.

However, even with organism and habitat variables, the study results remain.

Pheromone lures designed to aggregate the Douglas-fir beetle trap a multi-

157 species, non-target, flying beetle assemblage. The results indicate a potential community effect (of an unknown extent) resulting from pheromone baiting that may influence management applications. However, any applications will

require a firm understanding of the limitations of this and future studies.

Study limitations

Despite the significant and theoretically supported results of this study, four major aspects of study design impose limitations to study results. Addressing trapping bias, issues of pheromone composition, improving sampling design associated with replication and site selection, and addressing issues of spatial and temporal scale could clarify study results.

This study is limited by the use of a single trap design for species sampling.

Although the selection of the Lindgren trap was determined to be the best choice for the study, the interpretation of capture data obtained by a single technique may be incomplete, biased, or misleading (Muirhead-Thomson

1991). The pheromone bias measured in this study is significant and quantifiable through the use of control traps (investigating the extent of pheromone bias would not have been possible without the use of comparable, unbaited traps). What is not known however is how the observed species complement from either baited or control Lindgren traps would compare to trapping by "traditional" flight interception traps. Pheromone trapping influences the sampling diversity of flying beetles from preharvest and post

158 harvest conditions, but trapping also measures species potentially influenced by trap design, and secondary visual cues. Further research is needed to establish the impact of physical aspects of pheromone trapping (trap design, trap conditions, random interception) and how these elements compare to established passive trapping techniques. The results of such a comparison would not only create futher context to the bias of Lindgren trapping (both with and without pheromones), it would connect pheromone-biased sampling with other standard and comparable 'sampling packages' (Kitching et al. 2001) used in traditional survey methods. Such a comparative study would also confirm, refute, and further describe the species assemblages and patterns observed in this study.

Pheromone composition

As noted in the the methodology, the lure used in this study consisted of a ternary blend of racemic i1) frontalin (Fn) (1,5,-dimethyl-6,8- dioxabicyclo[3.2.1]octane), racemic i1) MCOL (1-methylcyclohex-2-enol) and seudenol (3-methylcyclohex-2-en-1-ol) of an undetermined enantiomeric composition. The design of the lure and resulting variability in semiochemical composition (see Chapter 2) may have impacted species responses. It is considered that elements of predictability and stability were present in semiochemical compostion and release, though the full extent of stability or varation is unknown and should be addressed in future studies.

159 The efficacy of pheromones warrants their continued use as a management tool, but the future development of operational pheromones may need to balance pheromone efficacy for the target species against the impact of pheromones on non-target species. A case in point is the use of ethanol, which is known to be synergistic in aggregating the target species (Pitman et al. 1975), but also attracts non-target wood-boring species (Mongomery and

Wargo 1983). When released as part of a pheromone lure ethanol is thought to increase trap catches of a number of non-target flying beetle species (Peck et al. 1997). The continued refinement of current lures coupled with increased efficacy and increased management applications, may warrant a greater assessment of the non-target impact of new pheromone components

(particularly highly parsimonious semiochemicals) for use in IPM containment and monitoring programs.

Site selection & replication

Changes in flight behavior are affected, or determined by a variety of subtle ecological, physiological, and meterological conditions (Muirhead-Thomson

1991). Site selection for this study was based on a narrow criterion for one dominant plant species. The presence and extent of Douglas-fir was thought appropriate given the study's focus on beetle-attacked Douglas-fir. However, with the large number of species ultimately gathered by the study, and the variation in species compostion and abundance observed between sites within and between harvest conditions/years, it may have been prudent to consider

160 the influence of site quality (Chapman 1955, Anonymous 1998) as it is known to influence species presence (Nilsson et al. 1994, Nilsson and

Baranowski 1997, Werner and Raffa 2000, Church et al. 2000). Differences in structural diversity, microhabitats, secondary plant species, and the amount of coarse woody debris in post harvest conditions, were unaccounted for in site selection and may have complicated study results.

An irreconcilable concern in this study involves sampling design associated with replication. The study utilized every available site that conformed to study criteria, and site availability was predetermined according to harvesting schedules, forest development plans, and accessibility. While none of these predisposing agents are thought to influence the results of this study, a truly random site selection of a sampling pool of all beetle attacked Douglas-fir stands in the District was not available.

The predetermined nature of site selection also created an uneven distribution of sampling sites for baited and control data across study years and biogeoclimatic subzones, predisposing the data to a systematic bias in spatial, temporal, and habitat conditions, both within and between treatments. Spatial bias occurred when preharvest and postharvest sampling was repeated in some sites, but not others (Table 3.1). Temporal bias occurred when sites respresenting a given treatment condition (ie. Preharvest conditions) were sampled nonsystematically across study years (see Table 2.1). Finally, a

161 habitat bias is present with the uneven representation of different biogeoclimatic zubzones in the study. The impact of these biases are unknown, however, the impact of such sampling bias' did not appear to be large enough to overwhelm the data or interfere with this assessment of the

Douglas-fir beetle pheromone lure.

Lastly, it is considered that the scale of the study must relate and apply to the question (Wiens et al. 1986), and issues of spatial and temporal scale (with respect to the context of study results), needs to be addressed. The Fort St

James Study is thought to be a medium (albeit on the small side of medium) field experiment. As a medium term field experiment in multiple sites the study should allow insight into causal factors determining the dynamics of the system (Wiens et al. 1986). The results of the study appear to approach this expectation by indicating a multispecies response to pheromones, and suggesting potential guild/habitat associations for those species based on observed abundance trends. Regrettably, the study is too broad to elucidate the nature of species responses, and too small to allow more than limited contextual generalizations about the nature of the community responding to pheromones. With regards to temporal scale, the study identifies short-term impacts on species diversity associated with a pheromone trapping-harvesting

IPM program for Douglas-fir beetle, but long-term impacts remain unknown.

Summary

162 The analysis of disturbance characteristics, including insect activity, is important to understanding ecosystem structure and function, and critical to effective resource management (Schowalter 1981, Lautenschlager 1997). In addition, insect responses to disturbance are thought to contribute to our views of community and ecosystem organization (Schowalter 1985, Miller

1993). The Douglas-fir beetle both creates and responds to disturbance events in Douglas-fir habitat, and is capable of influencing the rate and direction of forest succession. Its utilization of mature, overmature, and old- growth stands makes it an important species for integrated pest management

(IPM) programs. A simple three-semiochemical lure based on the Douglas-fir beetle pheromone system is highly effective in manipulating the distribution and abundance of natural Douglas-fir beetle populations as a tool for IPM programs, but results from this study indicate that pheromone lures influence the trap catch of non-target species in baited traps. The results may afford greater consideration of non-target impact resulting from pheromone applications, however the full extent of pheromone influence is unknown. The study results indicate short-term impacts on diversity associated with pheromone trapping and harvesting, but the long-term impacts are also unknown. Planning based on incomplete information on species as well as incomplete information on their existence and distribution creates a challenging problem (Church et al. 2000), and until further research is done to elucidate the impact of pheromone lures at the species level, the most readily

163 applicable management consideration may be to consider the scale of potential impact relative to habitat replacement.

Changes in trapped diversity resulting from pheromone lures were observed in addition to a dynamic and variable non-target multispecies assemblage present in endogenous and exogenous disturbance conditions associated with beetle attack, harvesting, and the onset of secondary succession. Trap catch data from management activity crossed compositional, structural, and functional levels of biodiversity, suggesting the potential for pheromone trapping to sampling flying beetle communities in a biased but concisely defined context.

Changing the distribution and abundance of multispecies assemblages impacts biodiversity. To preserve all species, in all locations under all circumstances, is logistically impossible (Van Kooten 1994). IPM programmes, by definition, go beyond a single objective of pest control to reflect a comprehensive approach to pest management - one that encompasses ecological, economic, and sociological impacts (Kogan and

Lattin 1993). Management means assessing the importance of what's being altered, assessing the scale of alteration relative to the ecosystem, and understanding how the alterations fit with both short and long term management objectives (Kangas and Kuusipalo 1993, Paulson 1995). The management of multispecies assemblages, whether by design or default,

164 needs to be better understood for effective long-term management (Maddock and Du Plessis 1999). In the context of sustainable forest management, this means going beyond a species-based approach to address the functional role(s) of the species/community involved. Taken in context the results of this study offer some insight into the functional organization of flying beetles in response to a combined pheromone + harvesting IPM program. The results also indicate a potential for pheromone-biased sampling to assess and monitor flying beetle communities associated with intensive management practices. It has been said that the interaction between one beetle and it's host cannot be studied in isolation (Birch 1984), and althought that statement appears to run contradictory to the efficiency of pheromone development, further research on the extent and nature of non-target pheromone influence could add a much needed context to the efforts of IPM programs, increasing the potential for effective management of beetle attacked, interior Douglas-fir.

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182 APPENDIX I Taxonomic Support

The following individuals generously supported this project with their time and expertise.

Family Taxonomic Specialist Affiliated Institution Alleculidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Anobiidae Don Bright Agriculture & Agrifood Canada, Ottawa Anthribidae Bob Skidmore Agriculture & Agrifood Canada, Ottawa Bostrichidae Bob Skidmore Agriculture & Agrifood Canada, Ottawa Buprestidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Byrrhidae Laurent LeSage Agriculture & Agrifood Canada, Ottawa Byturidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Cantharidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Carabidae Jeff Jarrod University of British Columbia, Canada Yves Bousquet Agriculture & Agrifood Canada, Ottawa Cerambycidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Cerylonidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Chrysomelidae Laurent LeSage Agriculture & Agrifood Canada, Ottawa Ciidae Don Bright Agriculture & Agrifood Canada, Ottawa Clambidae Anthony Davis Agriculture & Agrifood Canada, Ottawa Cleridae Bob Skidmore Agriculture 8c Agrifood Canada, Ottawa Coccinellidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Colydiidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Corylophidae Anthony Davis Agriculture & Agrifood Canada, Ottawa Cryptophagidae Richard Leschen Landcare Research New Zealand Ltd. Cucujidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Curculionidae Bob Anderson Canadian Museum of Nature, Ottawa Dermestidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Derodontidae Don Bright Agriculture & Agrifood Canada, Ottawa Dytiscidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Elateridae Ed Becker Agriculture & Agrifood Canada, Ottawa Erotylidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Eucnemidae Ed Becker Agriculture & Agrifood Canada, Ottawa Eucinetidae Don Bright Agriculture & Agrifood Canada, Ottawa Histeridae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Serge Laplante Agriculture & Agrifood Canada, Ottawa Hydraenidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Hydrophilidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Lampyridae Ale Smetana Agriculture & Agrifood Canada, Ottawa Lathridiidae Fred Andrews California Department of Food and Agriculture, Sacramento. Leiodidae Anthony Davis Agriculture & Agrifood Canada, Ottawa Lucanidae Serge Laplante Agriculture 8c Agrifood Canada, Ottawa Lycidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Melandryidae Darren Pollock University of Manitoba, Winnipeg Melyridae Don Bright Agriculture &. Agrifood Canada, Ottawa Mordellidae Don Bright Agriculture &. Agrifood Canada, Ottawa Mycetophagidae Yves Bousquet Agriculture &. Agrifood Canada, Ottawa

Family Taxonomic Specialist Affiliated Institution Nemonychidae Don Bright Agriculture & Agrifood Canada, Ottawa Nitidulidae Anthony Davies

183 Oedemerinae Serge Laplante Agriculture & Agrifood Canada, Ottawa Phalacridae Anthony Davies Agriculture & Agrifood Canada, Ottawa Pselaphidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Ptinidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Pyrochroidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Pythidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Rhizophagidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa Salpingidae Darren Pollock University of Manitoba, Winnipeg Scaphidiidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Scarabaeidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Scirtidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Scolytidae Don Bright Agriculture & Agrifood Canada, Ottawa Scraptiidae Darren Pollock University of Manitoba, Winnipeg Scydmaenidae Sean O'Keefe Moorhead State University, Kentucky Silphidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Sphaeritidae Anthony Davies Agriculture & Agrifood Canada, Ottawa Sphindidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Staphylinidae Ale Smetana Agriculture & Agrifood Canada, Ottawa Anthony Davis Agriculture & Agrifood Canada, Ottawa Stenotrachelidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Tenebrionidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Darren Pollock University of Manitoba, Winnepeg Tetratomidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Throscidae Serge Laplante Agriculture & Agrifood Canada, Ottawa Trogossitidae Yves Bousquet Agriculture & Agrifood Canada, Ottawa

184 APPENDIX II Procedures for Statistical Analysis of Douglas-fir Bark Beetle Trap Catches

Synthetic pheromones used in field research trials are designed to manipulate insect populations through olfactory responses. Seasonal totals of the numbers of beetles trapped synthetic pheromone blends in southeastern and central British Columbia routinely exceed 200,000 insects per season. Owing to the large number of beetles tapped every year, and accurate and time efficient method is desired for estimating the number of trapped beetles within a species. The use of weight as a ratio estimator is considered to be an option.

Beetle weight within a population, at a given time, is considered to be positively correlated, and proportional with beetle number. Weight is also considered to follow a normal distribution curve with a number of secondary factors influencing weight variation include feeding, water saturation, variable physiological factors (such as egg and sperm production), and post trapping desiccation, Using samples collected from single trapping seasons, correlations between beetle weight and number were obtained. For the sampling year 1993 an initial sample of 22 trap catches resulted in a correlation of 0.995. Regression analysis resulted in linear plots with equations passing near the origin.

y=85.65x+27.8

Weight measurements and beetle number were independent and dependent variable respectively. The observed deviation for origin intercept (thought to result from trapping and processing desiccation) will not adversely affect the use of the ratio estimation technique.

ESTIMATOR OF THE SAMPLE RATIO r:

r = ZXL where Xi = beetle weight of the i sample Syi yi = beetle number of the ith sample

ESTIMATED VARIANCE OF r:

V( r) = N-n • _1_ • U yi-iIXijL2

2 Nn Ltx n-1

BOUND ON THE ERROR OF ESTIMATION

1 t(V (r))'0. 5 t = critical t value [2.08 (n=21)] at 95% confidence.

185 For a sample n = 22 (out of a total of N=322), the following values were obtained:

E yi = 8893 I Xj = 9670

2 xi y, = 72763.58 u. = x = 4.4

r = 91.96 v(r) = 4.37

Bound = ± 2.08 (4.37)5

In general, r = y/x is a biased estimator of R = uy/ux however, the bias becomes negligible if the relationship between x and y is linear and runs through the origin. Due to the deviation from the origin, the bias of the ratio estimator is taken into consideration. A ratio bias of 0.0028 in this circumstance is not considered to be serious.

BIAS OF RATIO ESTIMATOR:

2 N - n • Sx - rho sy • s2 Nn x2 y x

ESTIMATION OF TOTAL BEETLE NUMBER FROM 1993 SAMPLES

E(Y) = Rx E(Y) = 91.96 x V(y) = x2 (V(r)) V(y) = x2(4.37)

Bound = + 1.96 (x2 (4.37))5

Where x is the weight of an unknown number of Douglas-fir beetles.

Results of the analysis indicate that ratio estimation techniques can be accurately used to estimate trap catch numbers of Douglas-fir beetles. Based on the above data the minimum number of samples required to achieve a maximum bound on the ratio of estimation of11.3 (~ + 5% of the observed r value) is calculated to be 75. To minimize and account for variation in the data (including feeding and seasonal physiological variation), 6 randomly chosen samples per week, through 14 weeks of collections should comprise the statistical analysis. This stratification of the data, in combination with a regimented sample cleaning and drying process, will allow for better representation of the natural and sampling induced variability throughout the season. Producing more accurate results.

186 APPENDIX III Species presentation by trends in abundance

Table 1 - Increasing abundance of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5* season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

L£5 CN m harves t 0 CO to to i o o o to Species - Increasing Abundance Family CL Q_ Q_ Q_ O CL Adalia bipunctata (Linneaus) Coccinellidae 0 0 0.2 0. 0.3 Aleochara sekanai Klimaszewski Staphylinidae 0 0.1 0 0. 0.2

Attica tombacina (Mannerheim) Chrysomelidae 0 0 0 1.2

Amara lunicollis Schodte Carabidae 0 0 0 0.4

Ampedus (nr) moerens (LeConte) Elateridae 0 1 0 0. 4.6

Ampedus moerens (LeConte) Elateridae 0 0 0 0. 0.6 Ampedus phoenicopterus Germar Elateridae 0 0 0.2 0. 0.2 Ampedus pullus Germar Elateridae 0 3.2 3 5.5

Ampedus pullus Germar Elateridae 0.1 2 1 6 Aphodius haemorrhoidalis (Linnaeus)/pectoralis LeConte Scarabaeidae 0.1 0.1 0.2 0. 1

Bembidion grapii Gyllenhal Carabidae 0 0 0 0. 0.4

Bolitopunctus muricatulus Staphylinidae 0.9 1 2 1. 2

Bradycellus lecontei Csiki Carabidae 0 0 0 0.4 Bradycellus neglectus (LeConte) Carabidae 0 0.4 0.5 0. 0.7 Bradycellus nigrinus (Dejean) Carabidae 0.1 0.5 1.1 2. 1.8

Bradycellus nigrinus (Dejean) Carabidae 0 0 0 1. 5.8 Bryophacis smetanai Staphylinidae 0 0.1 1 0. 1

Bryophacis smetanai Staphylinidae 0 0 0 0. 0.4 Buperstis langi Mannerheim Buprestidae 0 0 0 0. 0.3

Buperstis langi Mannerheim Buprestidae 0 0 0 0.4 Buprestis nutialli Kirby Buprestidae 0 0.2 0.7 0. 1 Byturus unicolor Say Byturidae 0 0 0.2 0. 0.3 Calitys scabra (Thunberg) Trogossitidae 0.4 1.3 2 4. 4.3 Callidium cicatricosum Mannerheim Cerambycidae 0 0.1 0 0. 0.2

Callidium cicatricosum Mannerheim Cerambycidae 0 0 0 0. 0.2

Carphonotus testaceus Casey Curculionidae 0 0 0 0. 0.2

Cercyon sp#2 (cinctus) Smetana Hydrophilidae 0 0 0 0.4 Coccinella septumpunctata Linneaus Coccinellidae 0 1 2.3 3. 2.3 Coccinella trifasciata perplexa Mulsant Coccinellidae 0 0.1 0.5 0. 1.5

Colon asperatum Leiodidae 0 0 0 0. 0.6 Cododera sp.#514 Cerambycidae 0 0 0.2 0. 0.2 Cosmosalia chrysocoma (Kirby) Cerambycidae 0 0.1 0.1 0. 0.2 Crepidodera sp Chrysomelidae 0 0 0.2 0. 0.2 Cryptophagus sp#3 Cryptophagidae 0.3 0.3 1 1. 1.5 Ctenicera aeripennis (Kirby) Elateridae 0.2 3.5 20 24. 19.3

187 Ctenicera aeripennis (Kirby) Elateridae 0 6 12 4. 14.4

Ctenicera angusticollis (Mannerheim) Elateridae 0.5 0.9 0.6 1.2

Ctenicera kendalli Kirby Elateridae 0 0 0 0. 0.4

Ctenicera nigricollis (Bland) Elateridae 1.5 2.5 3.7 3. 6.7 Ctenicera pudica(V\I.J. Brown +propola columbiana)(Lecon\e) Elateridae 5.5 9.3 10.4 27. 45.7 Ctenicera r. resplendens (Eschscholtz) Elateridae 0.2 3.8 11.6 18. 15.8 Ctenicera r. resplendens (Eschscholtz) Elateridae 0 4 4 13.8 Ctenicera semimetallica (Walker) Elateridae 0 0.2 0.4 0. 0.7 Ctenicera umbricola (Eschscholtz) Elateridae 0.7 11.6 18.8 14. 50.8 Ctenicera umbricola (Eschscholtz) Elateridae 0.4 4 6 24. 33.8 Cyphon concinnus (LeConte) Scirtidae 0 0 0 0.4 Cyphon sp(p) Scirtidae 0 0 0 2. 1

Dalopius (nr) tristis W.J. Brown Elateridae 0 0.5 1.5 1.7

Dalopius (nr) tristis W.J. Brown Elateridae 0 0 0 1. 2.6

Danosoma brevicorne (LeConte) Elateridae 0 2 0 3. 5.6

Dicerca tenebrica (Kirby) Buprestidae 0 1 3 1 8.8

Dichelonyx vicina (Fall) Scarabaeidae 0 0 0 0.5

Didion punctatum (Melsheimer) Coccinellidae 0 0 0 0.4

Epuraea flavomaculata Maklin Nitidulidae 0 0 0 0. 0.4

Eucnecosum tenue (LeConte) Staphylinidae 0 0.1 0.2 0. 0.2

Gabrius picipennis (Maklin) Staphylinidae 0 0 0 0. 1

Gyrophaena spp. Staphylinidae 0 0 0 0. 0.4

Hadrobregmus americanus (Fall) Anobiidae 0 0 0 0. 1.4

Hemicoelus carinatus (Say) Anobiidae 0 0.1 0.1 0. 0.2

Heterothops conformis Smetana Staphylinidae 0 0 0 0. 0.2

Hydnobius pumilus LeConte Leiodidae 0 0 1 0. 1

Hydnobius sp# 1 Leiodidae 0 0.5 0.7 0. 1.5 Hydnobius sp# 1 Leiodidae 0 0 0 0. 1 Hydnobius sp#2 Leiodidae 0 0.2 0.3 0. 0.7 Hydnobius sp # 2 Leiodidae 0 0 0 0. 0.8

Laccobius borealis Cheary /earn D.C. Miller Hydrophilidae 0 0 0 0. 0.4

Laemophloeus biguttatus (Say) Cucujidae 0 0 0 0. 1.5

Lelinohesperus borealis Staphylinidae 0 0 0 0.4

Leoides collaris (LeConte) Leiodidae 0 0.1 0 0. 0.2

Leoides collaris (LeConte) Leiodidae 0 0 0 0. 0.4 Leoides puncticollis/curvata C.G. Thomson/Mannerheim Leiodidae 0 0.2 0.3 0. 0.5

Leoides sp # 3 Leiodidae 0 0 0.1 0. 0.2

Leptusa sp Staphylinidae 0 0 0 0. 0.6

Limonius pectoralis LeConte Elateridae 0 0 0 0. 0.2

Lordithon cascadensis (Maikin) Staphylinidae 0 0 0 0. 0.3

Lordithon t. thoracicus (Fabricius) Staphylinidae 0 0.1 0.2 0. 0.7

Megasemum asperum (LeConte) Cerambycidae 0.1 0.3 0.8 0. 0.8

Neoclytus m. muricatulus (Kirby) Cerambycidae 0 0.1 0.1 0. 0.3

Pediacus depressus (Herbst) Cucujidae 0 0 0 0. 0.2

Phausis rhombica Fender Lampyridae 0 0 0 0. 0.2

Platycerus marginalis Casey Lucanidae 0 0 0 0.8

Ptinus californicus Pic Ptinidae 0 0 0 0. 0.2

Pygoleptura n. nigrella (Say) Cerambycidae 0 0 0 0. 0.8 Quedius m. molochinoides Smetana Staphylinidae 0 0 1 2. 1

188 Quedius rusticus/vilis Smetana Staphylinidae 0 0 0 0. 0.2

Scaphisoma castaneum Motschulsky Scaphidiidae 0 0 0 0. 0.2

Scolytus piceae (Swaine) Scolytidae 0 0 0 0. 0.2

Semijulistus ater (LeConte) Melyridae 0 0 2 2. 2

Sericus brunneus Elateridae 0 0 0.1 0. 1.3 sp#611 Scarabaeidae 0 0 0 0. 0.2

Stenichnus californicus Motschulsky Scydmaenidae 0 0.2 0.2 0. 0.3

Stenichnus californicus Motschulsky Scydmaenidae 0 0 0 0.4

Strictoleptura canadensis cribripennis (LeConteJ Cerambycidae 0.1 3.6 6.6 13. 8.3

Syntomus americanus (Dejean) Carabidae 0 0 0 0. 0.2

Tachinus thruppi Hatch Staphylinidae 0 0 0 0. 0.2

Tachyta rana (Casey/Say) Carabidae 0 0 0 0. 0.2

Teretruis montanus Horn Histeridae 0 0.2 0.4 1. 2.2

Tomoxia borealis (LeConte) Mordellidae 0 0 0 0. 0.2

Tribolium audax Halstead Tenebrionidae 0 0 1 0. 1.6

Trichodes ornatus hartwegianus A. White Cleridae 0 0 0 0.4

Trixagus sp Throscidae 0 0 0 0.4

Trixagus sp (carnicollis (Schaeffer)j Throscidae 0 0.1 0 0. 0.2

Tropideres fasciatus (Olivier) Anthribidae 0 0 0 0.3

189 Table 2 - decreasing abundance of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5* season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

in T— CM CO harve s to OO W to Species - Decreasing Abundance Family i— o O o o Q. Q_ Q_ CL 0. Abstrulia (nr) veriegatta Casey Tetratomidae 0.9 0.6 0.4 0.3 0.3 Agathidium depressum Fall /obtusum Hatch Leiodidae 3.7 3.5 3.1 0.9 1 Anaspis sp Scraptiidae 25.6 8.7 9.6 3.2 3.2

Aphodius leopardus Horn Scarabaeidae 0.7 0 0 0 0

Athous nigropilis Motschulsky Elateridae 0.3 0 0 0 0 Atrecus macrocephalus (Nordmann) Staphylinidae 0.6 0.3 0.3 0.2 0 Caenocara scymnoides LeConte Anobiidae 0.2 0 0 0 0

Cryptophagus sp#1 Cryptophagidae 0.3 0 0 0 0 Ctenicera hoppingi (Van Dyke) Elateridae 0.1 0.1 0 0 0 Cucujus claviceps Mannerheim Cucujidae 4 4 1.3 0.4 1 Dicentrus bluthneri LeConte Cerambycidae 0.2 0.2 0.1 0 0

Dicentrus bluthneri LeConte Cerambycidae 0.3 0 0 0 0

Dienopteroloma subcostatum (Maklin) Staphylinidae 0.3 0 0 0 0

Drasterius debilis LeConte Elateridae 3 4 1 0.6 0.4

Dryocetes caryi/schelti Hopkins/Swaine Scolytidae 0.2 0.2 0 0 0 Earota sp. Staphylinidae 1 1 0 0 0

Earota sp. Staphylinidae 0.3 0 0 0 0 Eleates explanatus Casey Tenebrionidae 0.3 0.2 0 0 0 Emmesa stacesmithi Hatch Melandryidae 0.4 0.1 0.1 0 0 Enicmus mendax Lathridiidae 0.1 0.1 0 0 0

Enicmus mendax Lathridiidae 0.3 0 0 0 0 Enicmus tenuicornis LeConte Lathridiidae 2.6 2.5 2.1 1.2 1.3 Eupraea terminalis Mannerheim Nitidulidae 0.2 0 0 0 0 Eusphalerum spp (mostly pothos (Mannerheim)) Staphylinidae 202 2.9 2.7 1.1 3.7

Hallomenus sp Scraptiidae 0.3 0 0 0 0 Lathridius n sp Lathridiidae 0.5 0.2 0 0 0

Lathridius n sp Lathridiidae 0.3 0 0 0 0 Lordithon fungicola Campbell Staphylinidae 1.2 0.4 0.2 0 0 Omalium spit 1 Staphylinidae 0.1 0.1 0 0 0 Omosita discoidea (Fabricius) Nitidulidae 3.5 1.3 0.6 0.1 0 Odhocis punctatus Casey Ciidae 1.4 0.4 0.5 0.3 0.2 Pelecomalium testaceum (Mannerheim) Staphylinidae 4.5 1.8 0.6 0.2 0.5 Phryganophilus collaris LeConte Melandryidae 0.1 0.1 0 0 0 Pidonia scripta (LeConte) Cerambycidae 0.5 0 0 0 0 Placusa tacomae Staphylinidae 0.1 0.1 0.1 0 0 Pytho sp#2 Pythidae 0.3 0.1 0.1 0 0 Quedius criddlei (Casey) Staphylinidae 0.4 0 0 0 0 Quedius plagiatus Mannerheim Staphylinidae 3.5 0.7 0.6 0.4 0.2 Rhizophagus pseudobrunneus Bousquet Rhizophagidae 0.2 0.2 0.2 0.1 0 Rhyzophagus dimidiatus Mannerheim Rhizophagidae 2.6 0.5 0.1 0.2 0.2

190 Salebius nr minax Cryptophagidae 0.6 0 0 0 0 Scierus annectans LeConte Scolytidae 14.9 9.6 4.3 0.6 0.8 sp# 10 Nitidulidae 0.1 0.1 0.1 0 0 Spondylis upiformis Mannerheim Cerambycidae 4.3 3.1 0.7 1 1.7 Stephostethus breviclavus (Fall) Lathridiidae 0.4 0.4 0.2 0.3 0 Stephostethus liratus (LeConte) Lathridiidae 0.9 0.5 0.2 0.5 0.2

Syneta pilosa W.J. Brown Chrysomelidae 0.3 0 0 0 0 Tachinus frigidus Erichson Staphylinidae 0.2 0.1 0.1 0 0 Tetratoma concolor LeConte Tetratomidae 1.9 0.2 0.3 0 0

Tetropium velutinum LeConte Cerambycidae 1.3 2 1 0 0 Thanasimus undatulus (Say) Cleridae 120 116 75.8 29.4 28.5 Thymalus marginicollis Chevrolet Trogossitidae 2.7 0.6 0.2 0 0

Thymalus marginicollis Chevrolat Trogossitidae 4.3 4 2 0 0 Trachysida a. aspera (LeConte) Cerambycidae 0.3 0.2 0.3 0 0 Trypodendron retusum (LeConte) Scolytidae 2 0.9 0.1 0.1 0.2 Trypodendron retusum (LeConte) Scolytidae 1.3 1 0 0 0 Trypodendron rufitarsis (Kirby) Scolytidae 0.7 0 0.1 0 0 Trypodendron rufitarsis (Kirby) Scolytidae 0.3 0 0 0 0 Xylechinus montanus Blackman Scolytidae 5.6 0.2 0.1 0 0 Xylechinus montanus Blackman Scolytidae 0.7 0 0 0 0

191 Table 3 - Increasing then decreasing abundance of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5* season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

£ in CD CN CO ?F Species with increasing then decreasing £Z CD to W to to u_ o O o o abundance Family Q_ CL CL CL CL sp # 328 Scarabaeidae 0.1 2 1 0 0

Abstrulia (nr) veriegatta Casey Tetratomidae 1.6 3 0 0 0 Acidota crenata (Fabricius) Staphylinidae 0.2 0.6 1.9 2.3 0.3 Acmaeops p. proteus (Kirby) Cerambycidae 0 0.4 0.5 0.2 0.3 Agathidium depressum Fall /obtusum Hatch Leiodidae 2.7 2 6 1 1.2 Agathidium difformis (LeConte) Leiodidae 0 0 0.2 0.1 0 Agathidium difformis (LeConte) Leiodidae 0.3 1 0 0 0 Agriotella occidentalis W.J. Brown Elateridae 0 0 0 0.4 0.2 Aleochara gracilicornis Bernhauer Staphylinidae 0 0.3 0.2 0.2 0.2 Aleochara rubricalis Casey Staphylinidae 0 0.1 0.5 0.1 0 Aleochara suffosa (Casey) Staphylinidae 0 0.9 0.4 0.1 0.2 Aleocharinae (misc.spp) Staphylinidae 1 2.6 0.9 0.1 0 Allandrus populi Pierce Anthribidae 0 0.1 0.1 0.1 0 Altica tombacina (Mannerheim) Chrysomelidae 0 0.2 0.6 0.3 0.2 Amara discors Kirby Carabidae 0 0 0.1 0.1 0 Amara erratica (Dutschmid) Carabidae 0 1 1 0.6 0.6 Amara familiaris (Dutschmid) Carabidae 0 0.1 0.1 0.1 0 Amara laevipennis Kirby Carabidae 0 0.4 0.6 0 0 Amara latior (Kirby) Carabidae 0 0 0.2 0 0 Amara littoralis Mannerheim Carabidae 0 0.3 0.5 1.2 0.3 Amara littoralis Mannerheim Carabidae 0 0 0 2.2 0 Amara lunicollis Schodte Carabidae 0 0.5 0.6 0.8 0.5 Amischa sp. Staphylinidae 0 0 0.2 0.3 0 Ampedus (nr) moerens (LeConte) Elateridae 0 4 8.4 1.7 3.7 Ampedus behrensi + phelpsi (Horn) Elateridae 0 2.9 14.5 4.2 7.2 Ampedus brevis (Van Dyke) Elateridae 0.7 3.5 5.5 4.3 4 Ampedus brevis (Van Dyke) Elateridae 2.4 6 5 3.4 3.6 Ampedus mixtus (miniipennis?) (Herbst) Elateridae 0 0.9 1.6 0.8 1.2 Ampedus moerens (LeConte) Elateridae 0 1.8 1.2 1.1 1.2 Ampedus nigrinus (Herbst) Elateridae 0.6 14.5 23.1 14 10.3 Ampedus nigrinus (Herbst) Elateridae 0.1 30 20 2.8 12.8 Ampedus occidentalis Lane Elateridae 0.2 3.5 5.6 3.7 3.8

Ampedus occidentalis Lane Elateridae 0 25 21 4.6 4.8 Anaspis sp#2 Scraptiidae 0 1.1 1.2 1.4 0.5 Anisotoma globososa Hatch Leiodidae 0 0.5 1.2 1.2 0.2 Anotylus rugosus (Farbricius) Staphylinidae 0 0.5 0.2 0.4 0 Anthaxia inomata (Randall) Buprestidae 0.2 1.7 0.5 0.3 0 Antherophagus sp#1 Cryptophagidae 0 4 0 0 0 Antherophagus sp#2 Cryptophagidae 0.1 0.5 0.8 0.1 0.2 Antherophagus sp#2 Cryptophagidae 0 0 0 0.4 0

192 Asemum striatum (Linneaus) Cerambycidae 0.1 5.1 0.3 0 0

Asemum striatum (Linneaus) Cerambycidae 0.1 8 0 0 0 Atheta dentata Staphylinidae 0.3 2.4 1.4 0.4 0 Athous rufiventris rufiventris (Eschscholtz) Elateridae 0.9 1.5 7.1 1 1.8 Athous rufiventris rufiventris (Eschscholtz) Elateridae 2.1 4 3 2.4 2.6 Atomaria sp #10 Cryptophagidae 0 0 0 0.3 0 Atomaria spit 12 Cryptophagidae 0.1 0.4 0.1 0.1 0

Atomaria sp # 12 Cryptophagidae 0 0 1 0.2 0 Atomaria sp#6 Cryptophagidae 0 0.1 0.1 0.1 0 Bembidion grapii Gyllenhal Carabidae 0 0.8 0.6 0.4 0 Bembidion nigripes (Kirby) Carabidae 0 0 0.1 0.1 0 Bembidion tetracolum Say Carabidae 0 0.1 0.4 0 0 Bembidion versicolor (LeConte) Carabidae 0 0 0.3 0.2 0 Bhyrrhus sp Byrrhidae 0 0.1 0.1 0 0 Bisnius picicornis Staphylinidae 0.3 1.7 2 0 0 Bius estriatus (LeConte) Tenebrionidae 0 1.5 0.3 0 0 Bradycellus congener (LeConte) Carabidae 0 0 0.2 0 0 Bryophacis arcticus Staphylinidae 0 0.2 0 0 0 Bryophacis punctulatus Staphylinidae 0.1 0.3 0.4 0.1 0

Bryophacis punctulatus Staphylinidae 0 1 1 0.2 0 Bryophacis spp Staphylinidae 0 0.1 0.2 0.2 0

Buprestis lyrata Casey Buprestidae 0 0 11 8.4 7.2 Caenocelis sp# 1 Cryptophagidae 0.3 2.3 1.6 0.6 0.5

Calitys scabra (Thunberg) Trogossitidae 0.4 0 5 1 1.8 Calopus angustus LeConte Oedemerinae 1 1.5 3 1 0.5 Calyptomerus oblongulus Mannerheim Clambidae 0 0.1 0.1 0 0 Canifa sp# 1 Melandryidae 0 0 0.2 0.3 0 Carphacis nepigonensis (Bernhauer) Staphylinidae 0.4 1.2 1.5 0.3 0.5

Carphacis nepigonensis (Bernhauer) Staphylinidae 0.1 1 2 0 0.4 Carphoborus vandykei Bruck Scolytidae 0 0.5 0.3 0.3 0 Cephaloon tenuicorne LeConte Cephaloidae 0.3 1.2 1.9 1.1 0.7 Cerycon sp#1 (herceus frigidus) Smetana Hydrophilidae 0 0.2 0.1 0.6 0 Cerylon castaneum Say Cerylonidae 0.6 0.8 1.3 0.9 0.7

Cerylon castaneum Say Cerylonidae 0.9 2 2 0.8 0.4 Ceutorhynchus punctiger Gyllenhal Curculionidae 0 0.1 0.1 0 0 Chrysobothris carinipennis LeConte Buprestidae 0 0 0.3 0.4 0

Chrysobothris carinipennis LeConte Buprestidae 0 0 0 0.6 0 Cis angustus Hatch Ciidae 0 0.2 0.6 0.2 0.3 Colon asperatum Leiodidae 0 0.4 0.4 0.4 0.3 Colon magnicolle Mannerheim Leiodidae 1.2 1.4 1.5 0.6 0.7 Corticaria n sp Lathridiidae 4.3 5.4 5.5 1.9 0.5 Corticeus praetermissus (Fall) Tenebrionidae 0.1 0.3 0.2 0 0 Corticeus subopacus (Wallis) Tenebrionidae 0.2 0.2 0.4 0.1 0

Corticeus subopacus (Wallis) Tenebrionidae 0.1 2 0 0 0

Corticeus tenuis (LeConte) Tenebrionidae 0 0 0 0.4 0 Cortodera m. militaris (LeConte) Cerambycidae 0.4 1.5 1.5 0.3 0.2

Cortodera sp. #514 Cerambycidae 0 0 5 0.4 0.2

Cosmosalia chrysocoma (Kirby) Cerambycidae 0 0 0 0.4 0 Cossonus pacificus Van Dyke Curculionidae 0.2 1.1 0.6 0.5 0.2 Cryphalus ruficollis Hopkins Scolytidae 0.1 0.7 0.1 0.2 0

193 Cryptophagus sp#4 Cryptophagidae 2.1 1 2 0.6 0.8 Ctenicera bombycina (Germar) Elateridae 0 0.3 0.4 0.6 0 Ctenicera lobata (Eschscholtz) Elateridae 0 0.1 0.1 0 0 Ctenicera lobata (Eschscholtz) Elateridae 0 0 0 0.4 0 Ctenicera lutescens (Fa\\)/sagitticollis (Eschscholtz) Elateridae 0.2 1.5 0.6 0.5 0.2 Ctenicera nebraskensis (Bland) Elateridae 0.4 5.6 8.8 9.6 5.2 Ctenicera nitidula (LeConte) Elateridae 0 0.1 0.4 0.3 0 Ctenicera triundulata (Randall) Elateridae 0 0 0.1 0.1 0 Ctenicera volitans (Eschscholtz) Elateridae 0.4 4 1 2.2 0.8 Cucujus claviceps Mannerheim Cucujidae 1.9 3 6 0.2 0 Cylistus coarctatus (LeConte) Histeridae 0 0.6 0.7 0 0 Cylistus coarctatus (LeConte) Histeridae 0 1 1 0 0 Cyphon sp(p) Scirtidae 0.1 0.3 1.1 1.5 0.7

Cytursa luggeri Leiodidae 0 0 0 0.4 0.2 Danosoma brevicorne (LeConte) Elateridae 0 1.7 3.3 5.6 3.8 Dendroctonus pseudotsugae Hopkins Scolytidae 7699 12859 10362 4152 1137

Dendroctonus pseudotsugae Hopkins Scolytidae 6.4 16 15 7.6 2 Dendroctonus rufipennis (Kirby) Scolytidae 0 0.1 0.2 0.1 0 Dendrophagus cygnaei Mannerheim Cucujidae 1.7 5 1.5 0.5 0.3 Dermestes talpinus Mannerheim Dermestidae 0.1 0.2 0.2 0 0 Dicerca tenebrica (Kirby) Buprestidae 0 2.2 9.4 9.6 7.5 Dicerca tenebrosa (Kirby) Buprestidae 0 3.3 3 0.9 1 Didion punctatum (Melsheimer) Coccinellidae 0 0.1 0.3 0.2 0.2 Dienopteroloma subcostatum (Maklin) Staphylinidae 0.1 0.5 0.1 0 0 Dolichocis manitoba Dury Ciidae 0.1 0.7 0.4 0.2 0.2 Drasterius debilis LeConte Elateridae 0.5 1.1 0.2 0.5 0 Dryocetes affaber (Mannerheim) Scolytidae 0.5 7.4 1.4 0.8 0.2 Dryocetes affaber (Mannerheim) Scolytidae 0.6 6 4 0.8 0.4 Dryocetes autographus (Ratzeburg) Scolytidae 1.1 18.8 5.9 2.1 0.7 Dryocetes autographus (Ratzeburg) Scolytidae 0.6 14 4 2 2.4 Dryocetes confusus Swaine Scolytidae 0.3 4.1 0.3 0 0 Dryocetes confusus Swaine Scolytidae 0.1 1 0 0 0 Dyctyopterus spp Lycidae 1 2 6 0.4 0.8 Elaphrus americanus Dejean Carabidae 0 0 0.1 0.1 0 Ellychnia corrusca (Linneaus) Lampyridae 0 0 0.3 0 0 Ellychnia corrusca (Linneaus) Lampyridae 0 0 1 0.2 0 Enoclerus nr. Scheaferi Barr Cleridae 0.1 0.4 0 0 0 Enoclerus nr. Scheaferi Barr Cleridae 0 3 0 0 0 Enoclerus sphegeus (Fabricus) Cleridae 0.2 1 0.5 0 0 Epiphanis sp # 338 Eucnemidae 0 0.1 0.1 0.1 0 Epuraea planulata Erichson Nitidulidae 0.1 1.5 1.5 0.3 0.2 Epuraea planulata Erichson Nitidulidae 0.3 3 3 0.2 0.4 Epuraea spit 1 Nitidulidae 0.2 3.1 0.9 0.3 0.2 Epuraea sp# 1 Nitidulidae 0.3 2 0 0.8 0.2 Eupraea truncatella Mannerheim Nitidulidae 0.3 2.1 1.4 0.1 0.2 Eupuraea spit 2 Nitidulidae 0 0 0.1 0.1 0

Evodinus monticola vancouveri Casey Cerambycidae 0.9 3 1 0 0 Gabrius picipennis (Maklin) Staphylinidae 0 2.4 5.8 2.4 1.2 Glischrochilus confluentus (Say) Nitidulidae 0.1 2.6 0.2 0.1 0.2 Glischrochilus quadrisignatus (Say) Nitidulidae 0 0.4 0.5 0 0

194 Gnathacmaeops pratensis (Laicharting) Cerambycidae 0 0.3 0.4 0.1 0

Gnathoncus barbatus Bosquet & Laplante Histeridae 0 0.5 0.2 0 0

Gnathoncus communis (Marseul) Histeridae 0 0 0.1 0.1 0

Gnathotrichus retusus LeConte Scolytidae 2.2 15.7 30.7 12.7 0.7

Gnathotrichus retusus LeConte Scolytidae 0.6 4 2 4.4 1.4

Grypeta sp. Staphylinidae 0 0.2 0 0 0

Gyrophaena spp. Staphylinidae 0 0.5 0.2 0.2 0

Hadrobregmus americanus (Fall) Anobiidae 0 0.3 0.5 1.1 0

Hadrobregmus quadrulus (LeConte) Anobiidae 0 0 0.2 0.1 0

Hallomenus sp Melandryidae 0.1 0.1 0.3 0 0

Hapalaraea dropephylla Staphylinidae 0 0.2 0 0 0

Hapalaraea megadhroides (Fauvel) Staphylinidae 0.1 1 0 0.1 0

Hapalaraea megadhroides (Fauvel) Staphylinidae 0.4 2 0 0 0

Hapalaraea sp #1 Staphylinidae 0.1 0.2 0 0 0

Harpalus animosus Casey Carabidae 0 0 0 1 0

Harpalus laevipes Carabidae 0 0.1 0.5 0 0

Harpalus somnulentus Dejean Carabidae 0 0.1 0.5 0.2 0

Harpalus somnulentus Dejean Carabidae 0 0 0 0.6 0.2 Helophorus orientalis Motscholsky/sempervarians

Angus Hydrophilidae 0 0.4 0.4 0.2 0.3

Henoticus sp# 1 Cryptophagidae 0.1 0.9 0.4 0 0

Heterothops conformis Smetana Staphylinidae 0 3.2 5.8 1.5 0.2

Heterothops sp Staphylinidae 0 0 0.3 0 0

Hoppingiana sp (hudsonica) (LeConte) Melyridae 0.2 0.2 1.1 0.3 0

Hoppingiana sp (hudsonica) (LeConte) Melyridae 0 0 0 0.4 0.2

Hydnobius pumilus LeConte Leiodidae 0 0.5 0.9 0.7 0.3

Hydraena sp (pacifica) Perkins Hydraenidae 0 0 0.2 0.2 0

Hydrobius fuscipes (Linneaus) Hydrophilidae 0 0.3 0.3 0.2 0

Hydrobius fuscipes (Linneaus) Hydrophilidae 0 0 0 0.6 0.2

Hydroporus sp# 1 Dytiscidae 0 0 0.1 0.1 0

Hydroporus sp#3 Dytiscidae 0 0.1 0.2 0.1 0

Hylastes nigrinus (Mannerheim) Scolytidae 13.2 140 120 44.2 8.2

Hylastes nigrinus (Mannerheim) Scolytidae 6.1 224 13 11.4 13.2

Hylastes ruber Swaine Scolytidae 5.1 54.2 34.5 9 10.3

Hylurgops porosus (LeConte) Scolytidae 0 105 20.4 3.5 1.7

Hylurgops porosus (LeConte) Scolytidae 0.1 82 2 0.2 0.6

Hylurgops rugipennis Mannerheim/Fitch Scolytidae 0.1 1 0 0 0

Hypnoidus bicolor (Eschscholtz) Elateridae 0 0 1 1.6 0.2

Ips perturbatus (Eichhoff) Scolytidae 0.1 5.3 0.8 0 0.2

Ips pedurbatus (Eichhoff) Scolytidae 0 4 2 0.6 0.2

Ips pini (Say) Scolytidae 0.1 1.7 0.5 0 0.5

Ips tridens (Mannerheim) Scolytidae 0 3.4 0.9 0.1 0.3

Judolia m. montivagens (Couper) Cerambycidae 0 0.4 2.5 0.7 0.3

Judolia m. montivagens (Couper) Cerambycidae 0.1 1 10 0.4 0.6

Laccobius borealis Cheary /card D.C. Miller Hydrophilidae 0 0 0 0.4 0

Lathridius ventralis Lathridiidae 0 0.2 0.1 0 0

Lathrobium negrum LeConte Staphylinidae 0 0.3 0 0 0

Leiodes strigata (LeConte) Leiodidae 0 4.8 1.4 0.5 0.7

Lelinohesperus borealis Staphylinidae 0 0.2 0.1 0.2 0 Leoides puncticollis C.G. Thomson /curvata

Mannerheim Leiodidae 0 0 1 0.6 0.2

195 Leoides rufipes (Gebler) Leiodidae 0 0 0.2 0.2 0

Leoides rufipes (Gebler) Leiodidae 0.1 1 1 0 0.8

Leptusa sp Staphylinidae 0.1 0.2 0.2 0.1 0

Limonius pectoralis LeConte Elateridae 0 0 0.2 0.2 0

Medon sp. (nr. Pallescens) Staphylinidae 0 0.1 0.4 0.1 0.3

Megadhrus angulicollis Staphylinidae 0.2 1.4 0.4 0.2 0

Megasemum asperum (LeConte) Cerambycidae 0.6 3 1 0.4 0.8

Megatoma sp (cylindrica) (Kirby) Dermestidae 0.6 3.7 0.9 0.7 0

Megatoma sp (cylindrica) (Kirby) Dermestidae 0.9 2 3 0 0

Megatoma verigatta (Horn) Dermestidae 1.1 2.1 1.8 0.8 0.2

Melandrya striata Say Melandryidae 0.1 0.1 0.4 0.2 0.3

Melanophila drummondi (Kirby) Buprestidae 0 0.5 0.4 0.2 0

Microbregma e. emarginatum (Duftschmid) Anobiidae 0.6 2.3 2.4 0.7 0

Microbregma e. emarginatum (Duftschmid) Anobiidae 0.7 1 6 0.4 0.2

Micropeplus smetanai Campbell Staphylinidae 0 0.5 0.1 0.2 0

Molamba obesa Casey Corylophidae 0 0.2 0 0 0

Monochamus spp Cerambycidae 0 1.4 0.9 0.3 0.3 Mycetophagus distinctus Hatch Mycetophagidae 0 0.6 0.5 0.3 0.3 Mycetoporus brunneus (Marsham) Staphylinidae 0 0 0.3 0 0 Mycetoporus rufohumoralis Staphylinidae 0.4 1 0.5 0.4 0.5 Myrmecocephalus arizonicus (Casey) Staphylinidae 0 0.1 0.1 0.1 0 Myrmecocephalus arizonicus (Casey) Staphylinidae 0 0 0 0.4 0 Myrmedophila americana (LeConte) Cryptophagidae 0 0.4 0.5 0.2 0.3 Neanthophlax mirificus (Bland) Cerambycidae 1.7 17 3 4.6 5 Negastrius tumescens LeConte Elateridae 1 29.1 10.6 3.4 5.8 Negastrius tumescens LeConte Elateridae 0 56 10 1 3.6 Neoclytus m. muricatulus (Kirby) Cerambycidae 0 0 0 0.6 0.2 Nitidotachinus tachyporus Staphylinidae 0 0 0.1 0.1 0 Nudobius cephalus (Say) Staphylinidae 0 3.1 1.8 0.9 0.3 Odontosphindus clavicornis Casey Sphindidae 0 0.3 1.2 0.5 0 Olophrum boreale (Paykull) Staphylinidae 0 0.1 0.1 0.2 0 Olophrum boreale (Paykull) Staphylinidae 0 0 0 0.4 0 Olophrum consimile (Gyllenhal) Staphylinidae 0 0 1 0.6 0.6 Omalium sp # 2 Staphylinidae 0 0.1 0.1 0 0 Omalium spp. Staphylinidae 0 0.3 0 0 0

Omosita discoidea (Fabricius) Nitidulidae 0.3 2 0 0 0

Orchesia (nr) castanea Scraptiidae 0.1 0.2 0.1 0.2 0

Orsodacne atra (Ahrens) Chrysomelidae 0.1 0.7 0.2 0.5 0.3

Odhotomicus caelatus (Eichhoff) Scolytidae 0.1 4 5.7 1.5 0.2

Odhotomicus caelatus (Eichhoff) Scolytidae 0 10 2 0.6 0.2

Ostoma ferrugina (Linneaus) Trogossitidae 0.3 1 1 0.4 0.4

Oxytelus sp. Staphylinidae 0 0.7 0 0 0

Pachyta lamed liturata Kirby Cerambycidae 0 1.9 0.4 0 0

Pactopus hornii (LeConte) Throscidae 0.4 0.6 1.1 0.3 0.7

Pactopus hornii (LeConte) Throscidae 0.1 2 2 0.2 1

Paromalus mancus Casey Histeridae 0.6 4.3 0.9 0.1 0.2

Pediacus depressus (Herbst) Cucujidae 0 0.3 0.4 0.2 0.2

Pediacus fuscus Erichson Cucujidae 0.2 4.6 9.2 1.4 1.5

Pediacus fuscus Erichson Cucujidae 0 4 4 1.8 0.8 Pelecomalium testaceum (Mannerheim) Staphylinidae 0.3 15 2 0.4 0.2

196 Phaleromela verigata Triplehorn Tenebrionidae 0 0.1 0.1 0 0 Philodrepa (?) Dropephylla sp (nrlongula) Staphylinidae 0.1 1.1 6.9 4.6 4.2 Philonthinii spp Staphylinidae 0.3 1.2 2 0.7 0.3 Philonthus furvus Nordmann Staphylinidae 0 0 0.1 0.1 0 Phloeophagus canadensis Van Dyke Curculionidae 0 0 0.2 0 0 Phloeotribus lecontei/picea Schedl/Swaine Scolytidae 0 1.5 0.3 0.2 0 Phyllotreta striolata (Fabricus) Chrysomelidae 0 0.1 0.1 0 0 Pissodes (nr) fiskei Hopkins Curculionidae 0 0.2 0 0 0 Pissodes fasciatus LeConte Curculionidae 0.2 1.6 0.5 0 0 Pissodes striatulus (Fabricus) Curculionidae 0 1.3 0.3 0 0

Pissodes striatulus (Fabricus) Curculionidae 0.1 7 0 0 0 Pissodes striatulus dubius Randall Curculionidae 0 0.5 0 0 0 Pityogenes hopkinsi Swaine Scolytidae 0.1 0.1 0.4 0.1 0.2 Pityogenes plagiatus (LeConte) Scolytidae 0.1 4.2 1 0.3 0 Pityokeines minutus (Swaine) Scolytidae 0 0.5 0.5 0.5 0 Pityokteines elegans Swaine Scolytidae 0 0 0.3 0 0 Pityophthorus nitidulus Swaine^ tuberculatus EschhoffJ Scolytidae 0.1 0.8 2.2 0.9 0.5 Pityophthorus nitidulus Swaine^ tuberculatus Eschhoff) Scolytidae 0.3 2 1 1.4 0.4 Pityophthorus opaculus LeConte Scolytidae 0 0.6 0.3 0.4 0.2

Pityophthorus opaculus LeConte Scolytidae 0 0 0 0.6 0 Pityophthorus pseudotsugae Swaine Scolytidae 0.1 3.2 0.5 0.2 0

Pityopthorus aquilus Blackman (+ aplanatus) Scolytidae 0 2 0 0 0 Platycerus marginalis Casey Lucanidae 0 0.1 0.7 0 0 Platysoma coarctatum LeConte Histeridae 0 0.2 0.1 0 0 Platysoma leconti Marseul Histeridae 0 0.1 0.5 0.5 0 Plesiocis sp Ciidae 0 0.3 0.1 0 0 Podabrus piniphilus (Eschscholtz) Cantharidae 0.1 0.4 0.5 0.1 0.3 Pogonocherus penicillatus LeConte Cerambycidae 0 1.5 0.3 0 0

Pogonocherus penicillatus LeConte Cerambycidae 0 11 0 0 0

Poliaenus oregonus (LeConte) Cerambycidae 0 1 1 0 0 Polygraphus convexifrons Wood Scolytidae 0.4 1 0.2 0.1 0 Polygraphus rufipennis (Kirby) Scolytidae 3.2 43.2 21.1 7.8 4.3

Polygraphus rufipennis (Kirby) Scolytidae 0.3 12 6 5.8 2.2 Priognathus monilicornis LeConte Pythidae 0 1.1 0.3 0.1 0.5 Pseudohadrotoma sp (perversa) (Fall) Dermestidae 0 0.5 0.5 0.1 0 Pseudohylesinus nebulosus LeConte Scolytidae 4.3 102 61.5 3.9 2.3 Psyllobora vigintimaculata (Say) Coccinellidae 0 0.2 0.1 0 0 Pterostichus adstrictus Eschscholtz Carabidae 0 0.1 0.1 0 0 Pytho sp # 1 Pythidae 0 0.1 0.3 0.2 0.2

Pytho sp # 1 Pythidae 0 1 1 0.2 0

Pytho spit 2 Pythidae 0.1 1 0 0 0 Quediini spp Staphylinidae 0 0.3 0.3 0..1 0

Quedius disticalios Staphylinidae 0 1 1 0 0 Quedius m. molochinoides Smetana Staphylinidae 0 0.5 1.1 1.3 1 Quedius rusticus/vilis Smetana Staphylinidae 0.3 0.7 2.8 0.3 0.5 Quedius transparens Motschulsky Staphylinidae 0 0 0.1 0.1 0 Quedius velox Smetana Staphylinidae 4.4 7.3 8.9 5.3 4.8 Quedius velox Smetana Staphylinidae 4.7 9 16 2.6 4.8 Rhagium inquisitor (Linneaus) Cerambycidae 0 9.7 10.4 1.9 0.3

197 Rhagium inquisitor (Linneaus) Cerambycidae 0 9 11 0.4 0.2

Rhizophagus pseudobrunneus Bousquet Rhizophagidae 0 0 0 0.4 0 Rhizophagus remotus LeConte Rhizophagidae 0.6 0.8 0.7 0.1 0.5 Rhyncolus brunneus Mannerheim Curculionidae 0.2 0.2 0.5 0.2 0.3 Rhyncolus macrops Buchanan Curculionidae 1.7 2.6 2.5 0.9 0.8

Rhyncolus macrops Buchanan Curculionidae 1.6 3 3 0.2 1.2

Rhyzophagus dimidiatus Mannerheim Rhizophagidae 0.1 1 1 0 0 Sacium lugubre LeConte Corylophidae 0 4.5 1.3 1.6 0

Scierus annectans LeConte Scolytidae 4.4 12 2 0.8 0 Scierus pubescens Swaine Scolytidae 0.1 0.2 0 0 0 Scolytus piceae (Swaine) Scolytidae 0 4.7 0.6 0.3 0.3 Scolytus sp (unispinosus) LeConte Scolytidae 0 15.7 14.2 1.2 0.2 Scolytus tsugae (Swaine) Scolytidae 1.3 9.2 1 0.1 0

Scolytus tsugae (Swaine) Scolytidae 0 7 0 0 0 Scolytus unispinosus LeConte Scolytidae 0.4 28 12.5 1.4 0.2 Scotochroa basalis LeConte Melandryidae 0.1 0.4 0.7 0.5 0 Sericoda quadripunctata (DeGeer) Carabidae 0.1 1.2 0.5 0.6 0 Serralopalpus substriatus Haldeman Melandryidae 0.6 1.5 0.7 0.1 0

Serralopalpus substriatus Haldeman Melandryidae 0.6 2 0 0 0 Siagonium stacesmithi Hatch Staphylinidae 0.6 0.8 2.1 1 0.2

Siagonium stacesmithi Hatch Staphylinidae 0 3 3 0.4 0.2 sp#1 Chrysomelidae 0 0.1 0.1 0 0 sp#11 Nitidulidae 0 0.1 0.1 0 0 sp#12 Nitidulidae 0 0.3 0.1 0 0 sp#13 Nitidulidae 0 0.5 0.1 0 0 sp#15 Nitidulidae 0 0.2 0.2 0 0 sp#3 Nitidulidae 0 0.1 0.1 0.1 0 sp#6 Nitidulidae 0 0.2 0 0 0 sp # 608 Scarabaeidae 0 0 0.1 0.1 0 Sphaeriestes alternatus (LeConte) Salpingidae 0 0.4 0.1 0.2 0 Sphaerites politus Duftschmid Spaeritidae 0 0 0.1 0.1 0 Stenus bilineatus J. Sahlberg Staphylinidae 0 0.8 0.3 0.2 0.2 Stephanopachys substriatus (Paykull) Bostrichidae 0 0.2 0.1 0 0

Strictoleptura canadensis cribripennis (LeConte) Cerambycidae 0 0 1 12.6 5.6 Syneta albida LeConte Chrysomelidae 0.3 1.3 1.3 0 0.2 Syneta pilosa W.J. Brown Chrysomelidae 0 0.6 0.4 0 0 Syntomus americanus (Dejean) Carabidae 0.1 0.6 0.5 0.5 0.3 Tachinus elongatus Gyllenhal Staphylinidae 0.1 0.5 0 0 0

Tachinus elongatus Gyllenhal Staphylinidae 0.3 2 1 0.2 0

Teretruis montanus Horn Histeridae 0 0 0 0.8 0.2 Tetropium velutinum LeConte Cerambycidae 0.4 1.5 0.6 0.1 0 Thalycra mixta H. Howden Nitidulidae 0 0.4 0.5 0 0

Thanasimus undatulus (Say) Cleridae 2 44 5 0.6 0.4' Thanatophilus lapponicus (Herbst) Silphidae 0 0.4 0.2 0.2 0 Tomoxia borealis (LeConte) Mordellidae 0 0.1 0.3 0.2 0.2 Trachypachus holmbergi Mannerheim Carabidae 0 2.2 1.3 0.1 0.2

Trachysida a. aspera (LeConte) Cerambycidae 0.3 1 3 0 0.4 Triadhron lecontei Horn Leiodidae 0 0.1 0.1 0 0 Tribolium audax Halstead Tenebrionidae 0.3 4.3 1.5 1.7 1 Trichocellus cognatus (Gyllenhal) Carabidae 0 1.8 1.9 2.1 1.3

198 Trichocellus cognatus (Gyllenhal) Carabidae 0 0 2 0.6 0.6

Trichodes ornatus hartwegianus A. White Cleridae 0 0 0.1 0.2 0

Trichophya pilicornis (Gyllenhal) Staphylinidae 0 1.2 0.6 0.2 0

Trypodendron Linneatum (Olivier) Scolytidae 1274 4013 485 3.4 1.3

Trypodendron Linneatum (Olivier) Scolytidae 13 598 2 1 0.2

Typhaea stercorea (Linneaus) Mycetophagidae 0 0.2 0 0 0

Xestoleptura tibialis (LeConte) Cerambycidae 0 0.2 0.8 0.2 0.2

Xyleta laevigata (Hellenius) Melandryidae 2.9 87.7 27.1 8.4 6

Xyleta laevigata (Hellenius) Melandryidae 0.4 82 70 6.2 7.4

Xyletinus rotundicollis R.E. White Anobiidae 0 0 0.2 0.2 0

Xylotrechus longitarsis (Casey) /undulatus (Say) Cerambycidae 0 0.5 1.1 0.6 0.8

199 Table 4 - Decreasing then increasing abundance of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5* season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

Species - Decreasing Then Increasing CO •9 sharve s w w (/) y) Abundance Family o o o o i CL CL CL CL CL Acidota• crenata (Fabricius) Staphylinidae 0.3 0 0 0. 1.2

Bolitopunctus muricatulus Staphylinidae 3 1 0 2. 2

Calopus angustus LeConte Oedemerinae 0.7 2 0 1.4

Corcodera (prob) longicornis (Kirby) Cerambycidae 0.1 0 0 0.2

Codicaria n sp Lathridiidae 2.6 1 1 1.8 Cryptophagus sp# 1 Cryptophagidae 0.4 0.4 0.1 0.3

Cryptophagus sp#3 Cryptophagidae 0.7 0 0 0. 0.8

Ctenicera angusticollis (Mannerheim) Elateridae 0.9 0 0 0. 1.2

Ctenicera mendax (LeConte) Elateridae 0.4 0 0 0.2

Ctenicera semimetallica (Walker) Elateridae 0.1 0 0 0. 1 Epuraea flavomaculata Maklin Nitidulidae 0.1 0.1 0 0.7

Eusphalerum spp (mostly pothos (Mannerheim)) Staphylinidae 6.9 6 1 3. 2.2 Evodinus monticola vancouveri Casey Cerambycidae 2.7 0.2 0 0. 0.2

Hydnobius sp#3 Leiodidae 0.1 0 0 0. 0.2

Lordithon cascadensis (Malkin) Staphylinidae 0.1 0 0 0. 0.6

Mycetoporus americanus Erichson Staphylinidae 0.1 0 0 0. 0.2

Mycetoporus rufohumoralis Staphylinidae 1.9 1 0 0. 0.6

Odontosphindus clavicornis Casey Sphindidae 0.1 0 0 0. 0.8

Philonthinii spp Staphylinidae 0.3 0 0 0. 0.4 Rhinosimus viridiaeneus Randall Salpingidae 2.1 1.8 0.7 1

Tachinus basalis Erichson Staphylinidae 0.7 0 0 0. 0.4

Tachinus frigidus Erichson Staphylinidae 0.1 0 0 0.2

Tetratoma concolor LeConte Tetratomidae 1.4 0 0 0.2 Trichochrous albedensis Blaisdell Melyridae 0.4 0.2 0.5 0. 0.7

Trichochrous albedensis Blaisdell Melyridae 2.9 0 0 0.4 Triplax dissimulator (Crotch) Erotylidae 0.1 0.1 0 0.2

200 Table 5 - No abundance Trend of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5* season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

co CD C£D .C CM CO to to to to Species - No Abundance Trend Family CD o o o O Q_ D. 0. D. a. Acmaeops p. protects (Kirby) Cerambycidae 0 1 0 0 0.2 Agathidium basalis Leiodidae 0 0 0.1 0 0.2 Agathidium spp Leiodidae 0.3 0.8 1 0.7 1.5

Agathidium spp Leiodidae 0.4 0 0 0.6 0.2 Aleochara castaneipennis Mannerheim Staphylinidae 0.1 0 0.2 0.2 0

Aleocharinae (misc.spp) Staphylinidae 0 1 0 0 0.2 Amara erratica (Dutschmid) Carabidae 0 1.2 0.5 1 0.2 Amara idahoana (Casey) Carabidae 0.1 0 0.1 0.1 0.2

Ampedus behrensi + phelpsi (Horn) Elateridae 0 2 2 1 4.2

Ampedus mixtus (miniipennis?) (Herbst) Elateridae 0.3 3 0 0.4 0.6

Ampedus phoenicopterus Germar Elateridae 0 1 0 0.8 0

Anaspis sp Scraptiidae 1.4 3 61 4 3.6

Anaspis sp#2 Scraptiidae 0 1 3 0 0.4

Anisotoma globososa Hatch Leiodidae 0.4 0 1 1.2 0.4 Anotylus tetracarinatus (Block) Staphylinidae 0 0 0.3 0 0.2

Anthaxia inornata (Randall) Buprestidae 0.1 0 1 0.2 0 Antherophagus sp# 1 Cryptophagidae 0.4 0.2 0.4 0.1 0.2 Aphodius fimetarius (Linneaus) Scarabaeidae 0.8 0.1 0 0.1 0 Aphodius leopardus Horn Scarabaeidae 0 0 0.3 0 0.2

Atheta dentata Staphylinidae 0.1 3 0 0 0.2 Athous nigropilis Motschulsky Elateridae 0.4 0 0.1 0 0 Atomairia sp# 13 Cryptophagidae 0 0 0.2 0 0.2 Atomaria sp#3 Cryptophagidae 0 0.2 0 0.1 0 Atrecus quadripennis (Casey) Staphylinidae 0.1 0 0 0.1 0 Bius estriatus (LeConte) Tenebrionidae 0.3 0 0 0.2 0 Bradycellus lecontei Csiki Carabidae 0 0.3 0.2 0.8 0.3 Buprestis lyrata Casey Buprestidae 0 6.5 4.5 4.5 6.2

Buprestis nuttalli Kirby Buprestidae 0 0 0 0 0.6

Byturus unicolor Say Byturidae 0 1 0 0.2 0

Caenocelis sp # 1 Cryptophagidae 0.3 0 2 0.6 1 Calathus advena (Leconte) Carabidae 0.1 0 0.3 0 0.2

Cephaloon tenuicorne LeConte Cephaloidae 0.6 0 2 0.2 1 Cis sp (fuscipes) Mellie Ciidae 0.3 0.3 0 0.3 0 Clavilispinus rufescens (Hatch) Staphylinidae 0 0.1 0 0.1 0

Coccinella septumpunctata Linneaus Coccinellidae 0 0 4 5.8 3.2

Coccinella trifasciata perplexa Mulsant Coccinellidae 0 1 1 0.8 2

Colon magnicolle Mannerheim Leiodidae 0 0 1 0 0.4 Corcodera (prob) longicornis (Kirby) Cerambycidae 0 0.2 0.1 0.3 0.2

Codiceus praetermissus (Fall) Tenebrionidae 0 1 0 0 0.2 Codiceus tenuis (LeConte) Tenebrionidae 0 0.1 0 0.1 0

201 Cortodera m. militaris (LeConte) Cerambycidae 0 4 0 0 0.6 Cossonus pacificus Van Dyke Curculionidae 0 2 0 0.2 0 Cryptophagus sp#2 Cryptophagidae 0.4 0.2 0.2 0.2 0.2 Cryptophagus sp#4 Cryptophagidae 0.7 0.6 0.8 0.5 0.2

Ctenicera bombycina (Germar) Elateridae 0.1 0 0 1 0.2

Ctenicera comes (W.J. Brown) Elateridae 0.1 0 1 0 0 Ctenicera crestonensis (W.J. Brown) Elateridae 0 0.5 0 0.3 0.3 Ctenicera kendalli Kirby Elateridae 0 0.2 0.4 0.1 0.7 Ctenicera lutescens (Fa\\)/sagitticollis (Eschscholtz) Elateridae 0.3 1 0 0.2 0 Ctenicera nebraskensis (Bland) Elateridae 1.4 1 0 5.2 24.6

Ctenicera nigricollis (Bland) Elateridae 1.7 8 0 1.8 10.2

Ctenicera nitidula (LeConte) Elateridae 0 2 0 0.6 0.2 Ctenicera pudica(\NJ. Brown)+propola columbiana (Leconte) Elateridae 5.1 43 1 20.2 28.6 Ctenicera volitans (Eschscholtz) Elateridae 2.6 1.9 1 2.3 1.3 Curimopsis sp Byrrhidae 0.1 0 0.1 0 0.2 Cyphon concinnus (LeConte) Scirtidae 0 0 0.2 0 0.2 Cydusa sp (subtestacea) (Gyllenhal) Leiodidae 0 0 0.1 0 0.3 Cytilus sp (alternatus) (Say) Byrrhidae 0 0.3 0.5 0.2 0.7 Cytilus sp (alternatus) (Say) Byrrhidae 0 1 0 0 0.2 Dendroctonus rufipennis (Kirby) Scolytidae 0.6 0 1 0.4 0 Dendrophagus cygnaei Mannerheim Cucujidae 0.3 0 2 0.2 1 Dermestes talpinus Mannerheim Dermestidae 0.1 0 0 0.2 0

Dicerca tenebrosa (Kirby) Buprestidae 0 5 0 1.6 0.4

Dolichocis manitoba Dury Ciidae 0 0 1 0 0.2 Dorcatoma (prob) americana # 87 Anobiidae 0.4 0 0.1 0 0 Dyctyopterus spp Lycidae 2.3 0.7 2.4 1.2 0.7

Eanus decoratus (Mannerheim) Elateridae 0 1 0 0.2 0

Eanus sp # 1 Elateridae 0.6 1 0 0.2 0

Enicmus tenuicornis LeConte Lathridiidae 2.7 0 3 1.8 1 Epiphanis cornutus Eschscholtz Eucnemidae 0.2 0 0.1 0 0 Epuraea (nr) populi Dodge Nitidulidae 0 0.2 0.1 0 0.2 Epuraea depressa Nitidulidae 1.1 0.1 0 0.2 0.2 Ernobius gentilis Fall Anobiidae 0 0.4 0 0 0.2 Ernobius gentilis Fall Anobiidae 0 1 0 0 0.6 Ernobius nigrans Fall Anobiidae 0 0.1 0 0.2 0

Eupraea truncatella Mannerheim Nitidulidae 0 2 0 0.2 0 Glischrochilus moratus W.J. Brown Nitidulidae 0 0.2 0.1 0 0.3 Grammoptera subargentata (Kirby) Cerambycidae 0 0.4 0.1 0.7 0.2

Grammoptera subargentata (Kirby) Cerambycidae 0.1 0 1 0.4 1.2 Harpalus fuscipalps Carabidae 0 0 0.2 0 0.2 Harpalus nigritarsis C.R. Sahlberg Carabidae 0 0.1 0 0.1 0

Hemicoelus carinatus (Say) Anobiidae 0.1 0 0 0.2 0 Hydnobius sp#3 Leiodidae 0 0.1 0 0.2 0 Hydroporus sp # 2 Dytiscidae 0 0 0 0.2 0 Hylastes longicollis Swaine Scolytidae 0.3 0.8 0.4 0.8 0.3

Hylastes longicollis Swaine Scolytidae 0.1 1 0 0 0.4

Hylastes ruber Swaine Scolytidae 7.3 26 1 3.2 7 Hylurgops reticulatus Wood Scolytidae 0 0.2 0 0.1 0 Hylurgops rugipennis Mannerheim/Fitch Scolytidae 0 0.8 0 0.1 0

202 Hypnoidus bicolor (Eschscholtz) Elateridae 0 0.5 0.7 0.2 0.5 Ips latidens (LeConte) Scolytidae 0 0.2 0 0.1 0

Ips pini (Say) Scolytidae 0 5 1 0 0.2

Ips tridens (Mannerheim) Scolytidae 0 2 1 0 0.2 Ischnosoma sp (fibratum) Staphylinidae 0 0.1 0 0.1 0 Lacon rorulentus (LeConte) Elateridae 0 2.4 0.9 0.4 2

Lacon rorulentus (LeConte) Elateridae 0 1 2 0 0.2 Lasconotus complex LeConte Colydiidae 0 0.2 0 0 0.2 Lasconotus intricatus Kraus Colydiidae 0.1 0.1 0 0.1 0

Leiodes strigata (LeConte) Leiodidae 0 1 0 0.2 0.2 Limonius aeger LeConte Elateridae 0 0.3 0.3 0.2 0.8

Limonius aeger LeConte Elateridae 0 1 0 0.2 0 Lordithon bimaculatus Staphylinidae 0.3 0 0 0.1 0

Lordithon fungicola Campbell Staphylinidae 0 0 1 0.2 0.4

Lordithon poecilus (Mannerheim) Staphylinidae 0 0.1 0.2 0 0.2

Lordithon t. thoracicus (Fabricius) Staphylinidae 0 1 0 0 0.2

Megadhrus angulicollis Staphylinidae 0.1 1 0 0.4 0

Megatoma verigatta (Horn) Dermestidae 0.7 13 0 0.2 0.2

Melanophila drummondi (Kirby) Buprestidae 0 1 0 0.4 0 Micropeplus laticollis Maklin Staphylinidae 0.1 0 0.3 0.1 0.2

Micropeplus laticollis Maklin Staphylinidae 0.1 0 0 0.2 0

Molamba obesa Casey Corylophidae 0.1 1 0 0 0.6

Monochamus spp Cerambycidae 0 4 0 0.2 0 Mulsantina picta (Randall) Coccinellidae 0.2 0.3 0.2 0.1 0.2 Mycetochara fraterna (Say) Tenebrionidae 0.1 0 0.3 0.1 0

Mycetophagus distinctus Hatch Mycetophagidae 0.4 0 0 1 0.4 Mycetoporus americanus Erichson Staphylinidae 0.1 0 0.3 0.3 0

Myrmedophila americana (LeConte) Cryptophagidae 0 0 1 0.2 0.8 Neanthophlax mirificus (Bland) Cerambycidae 0.3 2.8 1.4 1.4 5.7

Nudobius cephalus (Say) Staphylinidae 0 0 1 0.2 0.6 Olophrum consimile (Gyllenhal) Staphylinidae 0 0 0 0.2 0 Omalium sp (foraminosum MaklinJ Staphylinidae 0.2 0.1 0.1 0.2 0

Orchesia (nr) castanea Scraptiidae 0.1 0 0 0.4 0 Orphilis subnitidus LeConte Dermestidae 0 0.4 0.2 0.1 0.2

Orsodacne atra (Ahrens) Chrysomelidae 0.6 0 0 4.2 . 0.4

Orsodacne sp Chrysomelidae 0.1 0 0 0.4 0

Odhocis punctatus Casey Ciidae 0.3 0 2 0.2 0 Ostoma ferrugina (Linneaus) Trogossitidae 0.6 0.1 0.4 0.6 0.2 Oxytelus fuscipennis Mannerheim Staphylinidae 0.2 0 0.5 1.6 0.8

Oxytelus fuscipennis Mannerheim Staphylinidae 0.3 0 1 0.6 1.2

Paromalus mancus Casey Histeridae 0 4 6 0 0.2

Phausis rhombica Fender Lampyridae 0.1 0 0 0.2 0

Philodrepa (?) Dropephylla sp (nrlongula) Staphylinidae 0.4 2 4 1.6 4 Philonthus politus (Linneaus) Staphylinidae 0.1 0 0.1 0 0 Phymatodes dimidiatus (Kirby) Cerambycidae 0.5 0.6 0.5 0.2 0.5

Phymatodes dimidiatus (Kirby) Cerambycidae 0.1 0 1 0 0

Pissodes fasciatus LeConte Curculionidae 0.7 3 0 0.4 0.2 Pityogenes knetchteli Swaine Scolytidae 0 0.7 0 0.1 0

Pityogenes plagiatus (LeConte) Scolytidae 0 2 0 0.2 0.2

Pityophthorus pseudotsugae Swaine Scolytidae 0 2 0 0.2 0

203 Pityopthorus aquilus Blackman (+ aplanatus) Scolytidae 0 0.2 0.1 0.1 0.2 Platydema spit 1 Tenebrionidae 0 0 0 0.2 0 Plegaderus sayi Marseul Histeridae 0 0.1 0.1 0 0.2

Podabrus piniphilus (Eschscholtz) Cantharidae 0.6 3 0 0.4 0.6 Podabrus scaber LeConte Cantharidae 0 0 0.2 0 0.2

Podabrus scaber LeConte Cantharidae 0 2 0 0 0.2 Poliaenus oregonus (LeConte) Cerambycidae 0 0.2 0.1 0 0.2

Polygraphus convexifrons Wood Scolytidae 0.1 0 0 0.2 0

Pseudohylesinus nebulosus LeConte Scolytidae 4.7 647 0 1.6 1.2 Pseudopsis sp Staphylinidae 0.1 0 0.1 0.1 0 Ptinus californicus Pic Ptinidae 0 0.2 0.1 0.2 0.2 Pygoleptura n. nigrella (Say) Cerambycidae 0.1 0.6 0.5 0.9 0.7

Quediini spp Staphylinidae 0.1 0 0 0.4 0

Quedius plagiatus Mannerheim Staphylinidae 1.6 3 0 0.2 0 Quedius s. spelaeus Horn Staphylinidae 0.1 0 0.1 0 0

Rhinosimus viridiaeneus Randall Salpingidae 0.1 0 0 0 0.4

Rhizophagus remotus LeConte Rhizophagidae 0.6 0 0 0.4 0.2

Sacium lugubre LeConte Corylophidae 0.1 1 0 0.4 0.4 Salebius nr minax Cryptophagidae 0.4 0 0.2 0 0 Scaphisoma castaneum Motschulsky Scaphidiidae 0.1 0.4 0.3 0.9 0.7

Scolytus sp (unispinosus) LeConte Scolytidae 0.1 6 0 1.4 0.8

Scolytus unispinosus LeConte Scolytidae 0.1 5 0 1.6 1.6

Scotochroa basalis LeConte Melandryidae 0.6 1 0 0.6 0.2 Scymnus sp Coccinellidae 0 0 0.1 0 0.3

Semanotus ligneus (Casey) Cerambycidae 0 1 0 0 0.2 Semijulistus ater (LeConte) Melyridae 0 3.3 3.2 3.6 2.3 Silis d. difficilis LeConte Cantharidae 0.1 0.1 0 0.1 0 Sitona cylindricollis (Fahraeus) Curculionidae 0 0 0.1 0 0.2 sp#2 Ciidae 0 0.1 0 0.1 0 sp#3 Ciidae 0 0.3 0 0.1 0.2 sp#4 Nitidulidae 0 0.2 0 0 0.2 sp#9 Nitidulidae 0 0.4 0.1 0 0.3

Spondylis upiformis Mannerheim Cerambycidae 3.3 4 0 1 0.6 Staphylinus pleuralis LeConte Staphylinidae 0.1 0.5 0.7 0.5 0.7

Staphylinus pleuralis LeConte Staphylinidae 0.1 1 0 0.4 0.2 Stephostethus liratus (LeConte) Lathridiidae 0.4 0 0 1 0

Syneta albida LeConte Chrysomelidae 0.6 0 1 0.6 0 Syneta hamata Horn Chrysomelidae 0 0.1 0 0 0.7 Syntomium grahami Hatch Staphylinidae 0 0 0.2 0 0.2 Tachinus basalis Erichson Staphylinidae 0.7 0.1 0.1 0.1 0.3 Tachinus thruppi Hatch Staphylinidae 0.1 0 0.1 0.2 0 Tachyporus sp (canadensis Campbell) Staphylinidae 0 0 0.1 0 0.3

Thalycra mixta H. Howden Nitidulidae 0.4 0 0 0.2 0

Trachypachus holmbergi Mannerheim Carabidae 0 1 0 0.2 0 Tragosoma depsarium (Linneaus) Cerambycidae 0 0.1 0.3 0.1 0.3

Triadhron lecontei Horn Leiodidae 0.1 1 0 0 0.2 Trichophya pilicornis (Gyllenhal) Staphylinidae 0.1 1 0 0.6 0 Triplax antica LeConte Erotylidae 0 0.1 0 0 0.2 Triplax antica LeConte Erotylidae 0.1 0 0 0.2 0 Triplax californica LeConte Erotylidae 0.4 1 0.9 1.8 1.3

204 Triplax californica LeConte Erotylidae 0.7 1 0 0.8 1.2

Trixagus sp Throscidae 0 0.3 0 0.3 0.3

Trypodendron betulae Swaine Scolytidae 0 0 0 0.2 0

Trypophloeus populi Hopkins Scolytidae 0 0.2 0 1.4 0

Tychius picirostris (Fabricus) Curculionidae 0 0 0.1 0 0.2

Upis ceramboides (Linneaus) Tenebrionidae 0 0.2 0.1 0.2 0.2

Utobium elegans (Horn) Anobiidae 0.1 0 0.1 0.1 0

Xestoleptura tibialis (LeConte) Cerambycidae 0 1 0 0.4 0

Xylotrechus longitarsis (Casey) /undulatus (Say) Cerambycidae 0.1 0 1 1.2 0.4

Zilora occidentalis Mank Melandryidae 0.2 0 0.1 0 0.2

Zyras sp. Staphylinidae 0 0.3 0.3 0.3 0.2

205 Table 6 - Single occurrence species of flying beetles trapped in baited and unbaited Lindgren funnel traps from preharvest through 4/5 season postharvested Douglas-fir habitat (Fort St James Forest District, British Columbia). Baited funnels traps contained pheromone lures for the Douglas-fir beetle (Dendroctonus pseudotsugae) consisting of MCOL, seudenol, & frontalin.

Mean Abundance / Site

Single Occurrence Species Family preharves t Pos t 1 Pos t 2 Pos t 3 Pos t 4/ 5 sp # 296 Nitidulidae 0.1 0 0 0 0 sp # 299 Nitidulidae 0 0 0.1 0 0 Adalia bipunctata (Linneaus) Coccinellidae 0 0 0 0 0.2 Aegialia rufescens Horn Scarabaeidae 0 0 0.1 0 0 Agabus sp#619 Dytiscidae 0 0 0 0 0.2 Agabus sp # 622 Dytiscidae 0 0 0.1 0 0 Agathidium sp#1 Leiodidae 0 0 0.1 0 0 Aleochara (xeno) lanuginosa Gravenhorst Staphylinidae 0 0.1 0 0 0 Aleochara suffosa (Casey) Staphylinidae 0 0 0 0.2 0 Aleochara villosa Mannerheim Staphylinidae 0 0 0 0 0.2 Amara apricaria (Paykull) Carabidae 0 0.1 0 0 0 Amara apricaria (Paykull) Carabidae 0 0 0 0 0.2 Amara sinuosa (Casey) Carabidae 0 0.1 0 0 0 Anthobium reflexicolle Casey Staphylinidae 0 0 0 0.1 0 Anthrenus pimpinellae Fabricus Dermestidae 0 0 0.1 0 0 Aphodius distinctus (O.F. Muller) Scarabaeidae 0 0 0.1 0 0 Aphodius haemorrhoidalis/pectoralis (Linneaus)/LeConte Scarabaeidae 0 0 0 0 0.2 Aphodius opacus LeConte Scarabaeidae 0 0 0 0 0.2 Atomaria sp # 10 Cryptophagidae 0 0 1 0 0 Atomaria sp# 11 Cryptophagidae 0 0 0 0.1 0 Atomaria sp#4 Cryptophagidae 0 0.1 0 0 0 Atomaria sp#9 Cryptophagidae 0 0.1 0 0 0 Atrecus quadripennis (Casey) Staphylinidae 0 1 0 0 0 Attalus sp Melyridae 0 0.1 0 0 0 Bembidion canadianum Casey Carabidae 0 0 0 0 0.2 Bembidion quadrimaculatum (LeConte) Carabidae 0 0 0 0 0.2 Bembidion timidum (LeConte) Carabidae 0 0.1 0 0 0 Bhyrrhus sp Byrrhidae 0.1 0 0 0 0 Bledius ruficornis LeConte Staphylinidae 0 0.1 0 0 0 Bradycellus congener (LeConte) Carabidae 0 0 0 0 0.2 Bradycellus neglectus (LeConte) Carabidae 0 0 0 0 0.2 Bromius obscurus (Linneaus) Chrysomelidae 0 0 0 0.1 0 Bryophacis canadensis Staphylinidae 0.1 0 0 0 0 Bryophacis spp Staphylinidae 0 0 0 0 0.2 Calathus advena (Leconte) Carabidae 0.1 0 0 0 0 Canifa sp# 1 Melandryidae 0 0 0 0.2 0 Cardiophorus (prob) tenebrosus LeConte Elateridae 0 0 0 0.1 0 Carphoborus vandykei Bruck Scolytidae 0 0 0 0.2 0 Cercyon sp#3 (tolfino) Hatch Hydrophilidae 0 0 0 0 0.2

206 Cerycon sp#1 (herceus frigidus) Smetana Hydrophilidae 0 0 0 0.2 0 Ceutorhynchus erysimi (Fabricius) Curculionidae 0 0 0 0 0.2 Ceutorhynchus punctiger Gyllenhal Curculionidae 0 0 0 0 0.2 Cimberis (prob) turbans Nemonychidae 0 0 0.1 0 0 Cis angustus Hatch Ciidae 0 0 0 0 0.2 Cis sp (fuscipes) Mellie Ciidae 0 0 0 0.2 0 Clytus sp Cerambycidae 0 0 0 0.1 0 Colon (mylochus) aedeagosum Hatch Leiodidae 0 0.1 0 0 0 Colon (mylochus) aedeagosum Hatch Leiodidae 0.1 0 0 0 0 Colon sp # 1 Leiodidae 0 0 0 0 0.2 Colon sp# 1 Leiodidae 0 0 0 0.2 0 Colopterus truncatus (Randall) Nitidulidae 0 0 0.1 0 0 Corticaria sp Lathridiidae 0 0.1 0 0 0 Corticarina (prob) cavicollis (Mannerheim) Lathridiidae 0 0 0.1 0 0 Creophilus maxillosus (Linneaus) Staphylinidae 0 0 0.1 0 0 Crepidodera sp Chrysomelidae 0 0 0 0 0.2 Cryphalus ruficollis Hopkins Scolytidae 0 0 0 0 0.2 Cryptophagus sp # 2 Cryptophagidae 0 0 0 0.2 0 Ctenicera bipunctata (W.J. Brown) Elateridae 0 0 0 0 0.2 Ctenicera comes (W.J. Brown) Elateridae 0 0 0 0.1 0 Ctenicera sp-134 Elateridae 0 0.1 0 0 0 Curimopsis sp Byrrhidae 0 0 0 0 0.2 Cytursa luggeri Leiodidae 0 0 0 0 0.2 Dendroides ephemeroides (Mannerheim) Pyrochroidae 0 0 0.1 0 0 Dermestes lardarius Linneaus Dermestidae 0 0 0 0 0.2 Dermestes sp Dermestidae 0 0 0 0.1 0 Desmatogaster subconnata (Fall) Anobiidae 0 0 1 0 0 Dichelonyx vicina (Fall) Scarabaeidae 0 0 0 0 0.2 Diphyllcis ? sp Ciidae 0 0 0 0.1 0 Diplotaxis brevicollis LeConte Scarabaeidae 0 0 0 0.1 0 Diplotaxis brevicollis LeConte Scarabaeidae 0 0 0 0 0.2 Dorcatoma (prob) americana # 87 Anobiidae 0.1 0 0 0 0 Dryocetes betulae Hopkins Scolytidae 0.1 0 0 0 0 Dryocetes caryi/schelti Hopkins/Swaine Scolytidae 0 0 1 0 0 Eanus decoratus (Mannerheim) Elateridae 0 0 0.1 0 0 Elaphrus clairvillei Kirby Carabidae 0 0 0 0.1 0 Elaphrus clairvillei Kirby Carabidae 0 0 0 0 0.2 Eleates explanatus Casey Tenebrionidae 0.1 0 0 0 0 Enoclerus sphegeus (Fabricus) Cleridae 0 0 0 0.2 0 Epiphanis cornutus Eschscholtz Eucnemidae 0 0 0 0 0.2 Eucinetus (nr.) oviformis LeConte Eucinetidae 0 0.1 0 0 0 Eucnecosum tenue (LeConte) Staphylinidae 0 0 0 0 0.2 Eupraea terminalis Mannerheim Nitidulidae 0.1 0 0 0 0 Glischrochilus confluentus (Say) Nitidulidae 0 0 0 0 0.2 Glischrochilus moratus W.J. Brown Nitidulidae 0 0 0 0 0.2 Gymnusa atra Casey Staphylinidae 0 0 0.1 0 0 Gymnusa pseudovariegata Klimaszewski Staphylinidae 0 0 0.1 0 0 Gymnusa pseudovariegata Klimaszewski Staphylinidae 0 0 0 0.2 0 Gymnusa sp (grandiceps Casey) Staphylinidae 0 0 0.1 0 0 Gyrohypnus fracticornis (O.F. Muller) Staphylinidae 0 0 0.1 0 0

207 Hallomenus sp Scraptiidae 0 0.1 0 0 0 Harpalus obnixus Casey Carabidae 0 0 0.1 0 0 Harpalus opacipennis (Haldeman) Carabidae 0 0.1 0 0 0 Henoticus sp # 1 Cryptophagidae 0 1 0 0 0 Henotiderus lorna (Hatch) Cryptophagidae 0 0.1 0 0 0 Heterothops fraternus Smetana Staphylinidae 0 0.1 0 0 0 Hippodamia tredecimpunctata (Say) Coccinellidae 0 0.1 0 0 0 Hippuriphila sp Chrysomelidae 0 0 0 0 0.2 Hydaticus aruspex Clark Dytiscidae 0 0 0 0 0.2 Hygrotus impressopunctatus (Schaller) Dytiscidae 0 0 0 0.1 0 Hypnoides impressicollis (Mannerheim) Elateridae 0 0.1 0 0 0 Ips latidens (LeConte) Scolytidae 0.1 0 0 0 0 Ips mexicanus (Hopkins) Scolytidae 0 0.1 0 0 0 Isomera (nr) comstoki Papp Alleculidae 0 0 0.1 0 0 Laccobius sp Hydrophilidae 0 0 0 0.1 0 Laemophloeus biguttatus (Say) Cucujidae 0 0 0 0 0.2 Laricobius laticollis Fall Derodontidae 0 0.1 0 0 0 Lathridius ventralis Lathridiidae 0 0 0 0.2 0 Lathrobium negrum LeConte Staphylinidae 0.1 0 0 0 0 Lebia moesta LeConte Carabidae 0 0 0 0.2 0 Leoides sp#3 Leiodidae 0 0 0 0 0.2 Leoides sp # 49 Leiodidae 0 0.1 0 0 0 Macronaemia episcopalis (Kirby) Coccinellidae 0 0 0.1 0 0 Magdalis alutacea LeConte Curculionidae 0.1 0 0 0 0 Malthodes sp. Cantharidae 0 0 0 0 0.2 Malthodes sp. Cantharidae 0 0 0 0.2 0 Margarinotus rectus (Casey) Histeridae 0 0.1 0 0 0 Micetoporus sp Staphylinidae 0.1 0 0 0 0 Micetoporus sp Staphylinidae 0 0 0 0 0.2 Mycetochara fraterna (Say) Tenebrionidae 0 0 0 0.2 0 Mycetophagus tenuifasciatus Horn Mycetophagidae 0 0 0.1 0 0 Mycetophagus tenuifasciatus Horn Mycetophagidae 0 0 0 0 0.2 Mycetoporus brunneus (Marsham) Staphylinidae 0 0 0 0 0.2 Mycetoporus rugosus Hatch Staphylinidae 0 0 0 0.2 0 Myremcocephalus arizonicus (Casey) Staphylinidae 0 0.1 0 0 0 Neohypdonas tumescens (LeConte) Elateridae 0 0 0 0.1 0 Neohypnus obscurus (Erichson) Staphylinidae 0 0.1 0 0 0 Neohypnus obscurus (Erichson) Staphylinidae 0 0 0 0 0.2 Notiophilus aquaticus (Linneaus) Carabidae 0 0 0 0.1 0 Notiophilus directus Casey Carabidae 0 0 0.1 0 0 Ochthebius sp Hydraenidae 0 0.1 0 0 0 Ochthephilus planus (LeConte) Staphylinidae 0 0.1 0 0 0 Octotemnus denudatus Casey Ciidae 0 0 0.1 0 0 Octotemnus denudatus Casey Ciidae 0 0 0 0.2 0 Oiceoptoma noveboracense (Forster) Silphidae 0 0 0 0.1 0 Omalium n. sp. Staphylinidae 0 0 0.1 0 0 Omalium sp# 1 Staphylinidae 0 1 0 0 0 Omalium spp. Staphylinidae 0.1 0 0 0 0 Orchesia ornata Scraptiidae 0 0 0 0.1 0 Orsodacne sp Chrysomelidae 0 0 0 0 0.2

208 Orus sp Staphylinidae 0 0 0 0.1 0 Pachybrachis melanostictus Suffrain Chrysomelidae 0 0 0 0.1 0 Pachybrachis melanostictus Suffrain Chrysomelidae 0 0 0 0.2 0 Phaedon laevigatus (Duftschmid) Chrysomelidae 0 0 0 0.2 0 Philonthus concinnus (Gravenhorst) Staphylinidae 0 0.1 0 0 0 Philonthus couleensis Hatch Staphylinidae 0 0.1 0 0 0 Philonthus crotchi Horn Staphylinidae 0 0.1 0 0 0 Philonthus varians (Paykull) Staphylinidae 0 0.1 0 0 0 Phloeopra sp. Staphylinidae 0 0.1 0 0 0 Phloeosinus pini Swaine Scolytidae 0 0.1 0 0 0 Phloeotribus lecontei Schedl /picea Swaine Scolytidae 0.1 0 0 0 0 Phymatodes (nr) fulgidus Hopping Cerambycidae 0 0 1 0 0 Phymatodes maculicollis LeConte Cerambycidae 0.1 0 0 0 0 Pidonia scripta (LeConte) Cerambycidae 0.1 0 0 0 0 Pissodes (nr) fiskei Hopkins Curculionidae 0 1 0 0 0 Pissodes striatulus dubius Randall Curculionidae 0 1 0 0 0 Pityomacer pix Kuschel Nemonychidae 0 0.1 0 0 0 Pityopthorus sp Scolytidae 0 0.1 0 0 0 Plateumaris rufa (Say) Chrysomelidae 0 0 0 0.2 0 Platydema americanum Castelnau & Brulle Tenebrionidae 0 0.1 0 0 0 Platydema sp # 1 Tenebrionidae 0 1 0 0 0 Platysoma leconti Marseul Histeridae 0 0 0 0.2 0 Plegaderus setulosus Ross Histeridae 0 0.1 0 0 0 Podabrus fissilis Fall Cantharidae 0 0 0 0.1 0 Podabrus sp # 613 Cantharidae 0 0 0.1 0 0 Pogonocherus mixtus Haldeman Cerambycidae 0 0.1 0 0 0 Proteinus sp. Staphylinidae 0 0.1 0 0 0 Pselaphus bellax Casey Pselaphidae 0 0.1 0 0 0 Pseudohadrotoma sp (perversa) (Fall) Dermestidae 0 1 0 0 0 Psyllobora vigintimaculata (Say) Coccinellidae 0.1 0 0 0 0 Pterostichus adstrictus Eschscholtz Carabidae 0.1 0 0 0 0 Ptilinus lobatus/basalis Casey/Leconte Anobiidae 0 0 0 0 0.2 Quedius erythrogaster Mannerheim Staphylinidae 0.1 0 0 0 0 Quedius pediculus (Nordmann) Staphylinidae 0 0 0 0.1 0 Quedius transparens Motschulsky Staphylinidae 0 0 1 0 0 Rhantus binotatus (Harris) Dytiscidae 0 0.1 0 0 0 Rhyncolus brunneus Mannerheim Curculionidae 0 0 0 0 0.2 Saprinus lugens Erichson Histeridae 0 0.1 0 0 0 Scaphidema aeneolum (LeConte) Tenebrionidae 0 0 0.1 0 0 Scaphium sp Scaphidiidae 0 0 0 0.2 0 Scolytus opacus Blackman Scolytidae 0 0.1 0 0 0 Scolytus subscaber LeConte Scolytidae 0 0.1 0 0 0 Scraptiidae sp#3 Scraptiidae 0 0 0 0 0.2 Semanotus ligneus (Casey) Cerambycidae 0 0.1 0 0 0 Sepedophilus littoreus (Linneaus) Staphylinidae 0 0 0 0.2 0 Sericoda quadripunctata (DeGeer) Carabidae 0 0 0 0 0.2 Sericus brunneus Elateridae 0 1 0 0 0 Silis d. difficilis LeConte Cantharidae 0 0 0 0.2 0 Sitona lineellus (Bonsdorff) Curculionidae 0 0 0.1 0 0 Sitona lineellus (Bonsdorff) Curculionidae 0 0 0 0.2 0

209 Sonoma parviceps (Maklin) Staphylinidae 0 0 0 0 0.2 sp#1 Chysomelidae 0 0 0.1 0 0 sp#1 Anobiidae 0 0 0 0 0.1 sp#14 Nitidulidae 0 0.1 0 0 0 sp#2 Ciidae 0 0 0 0.2 0 sp#3 Nitidulidae 0 1 0 0 0 sp#3 Ciidae 0 0 0 0.2 0 sp#4 Nitidulidae 0 0 0 0.2 0 sp # 573 Throscidae 0 0 0 0 0.2 sp # 609 Scarabaeidae 0 0 0 0.1 0 sp # 610 Scarabaeidae 0 0.1 0 0 0 sp # 610 Scarabaeidae 0 0 0 0 0.2 sp#611 Scarabaeidae 0 0 0 0 0.2 sp # 621 Dytiscidae 0 0 0 0.2 0 sp # 623 Dytiscidae 0 0 0.1 0 0 sp # 624 Dytiscidae 0 0 0 0 0.2 sp # 625 Dytiscidae 0 0 0.1 0 0 sp # 626 Dytiscidae 0 0 0.1 0 0 sp#7 Nitidulidae 0.1 0 0 0 0 sp#8 Nitidulidae 0 0.1 0 0 0 sp#1 Phalacridae 0 0 0.1 0 0 sp#1 Phalacridae 0 0 0 0 0.2 Sphaeridium bipustulatum Fabricius Hydrophilidae 0 0 0.1 0 0 Sphaeriestes sp. Salpingidae 0 0 0.1 0 0 Stenolophus fuliginosus Dejean Carabidae 0 0.1 0 0 0 Stenotrachelus aeneus (Fabricius) Stenotrachelidae 0 0 0 0 0.2 Stenus juno Paykull Staphylinidae 0 0 0 0.1 0 Stenus plicipennis (Casey) Staphylinidae 0 0 0.1 0 0 Stephostethus cinnamopterus (Mannerheim) Lathridiidae 0 0.1 0 0 0 Syneta hamata Horn Chrysomelidae 0.1 0 0 0 0 Synuchus impunctatus (Say) Carabidae 0 0 0.1 0 0 Tachinus nigricornis Mannerheim Staphylinidae 0.1 0 0 0 0 Tachinus nigricornis Mannerheim Staphylinidae 0.1 0 0 0 0 Tachinus vergatus Campbell Staphylinidae 0 0 0 0.2 0 Tachyporus sp Staphylinidae 0 0 0 0.1 0 Tachyporus sp Staphylinidae 0.1 0 0 0 0 Tachyporus sp (lecontei CampbellJ Staphylinidae 0 0 0 0.1 0 Tachyta angulata Casey Carabidae 0 0 0 0 0.2 Thanatophilus lapponicus (Herbst) Silphidae 0 0 1 0 0 Tragosoma depsarium (Linneaus) Cerambycidae 0 0 0 0 0.2 Triplax dissimulator (Crotch) Erotylidae 0.1 0 0 0 0 Xestocis sp Ciidae 0 0 0 0 0.2 Zilora occidentalis Mank Melandryidae 0.1 0 0 0 0 Zyras sp. Staphylinidae 0 0 0 0 0.2

210 APPENDIX IV Tabular presentation of means and 90% confidence intervals for diversity indices presented in Chapter 3.

Table IV-1. Diversity as measured by Species Richness (S) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps were baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site.

Baited/ Preharvest Post 1 Post2 Post 3 Post 4/5 Control

Baited 65.00 + 15.46 1 24.80 + 23.25 1 45.40 + 25.67 98.80 + 16.40 97.6 + 6.34

Control 53.60 + 12.56 1 52.00 113.00 88.20 + 18.30 92.4 + 13.42

Table IV-2. Diversity as measured by Margalefs richness index (d) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site.

Baited/ Preharvest Post-| Post2 Post 3 Post 4/5 Control ,

Baited 7.02 + 1.54 13.84 + 2.02 15.03 + 2.19 1 2.37 + 2.04 1 3.39 + 1.77

Control 10.38 + 2.06 1 9.52 1 6.14 1 5.44 + 2.42 1 5.57 + 1.86

Table IV-3. Diversity as measured by Pielou's evenness index (J) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited / Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 0.13 + 0.05 0.13 + 0.04 0.12 + 0.03 0.18 + 0.05 0.38 + 0.16

Control 0.85 + 0.05 0.54 0.81 0.85 + 0.02 0.80 + 0.04

211 Table IV-4. Diversity as measured by Brillouin index from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited / Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 0.53 + 0.23 0.61 + 0.20 0.60 + 0.14 0.79+ 0.24 1.63+ 0.71

Control 2.95 + 0.28 2.04 3.53 3.39 + 0.16 3.26 + 0.12

Table IV-5. Diversity as measured by Shannon-Wiener index (H'i0) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5,h season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site.

^aitfd./ Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 0.23 + 0.10 0.28 + 0.09 0.27 + 0.06 0.36 + 0.11 0.75+0.34

Control 1.46 + 0.14 1.09 1.67 1.64 + 0.06 1.57 + 0.06

Table IV-6. Diversity as measured by Simpson's dominance index (1- ) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited / Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 0.24 + 0.16 0.17 + 0.07 0.19 + 0.07 0.23 + 0.07 0.51 + 0.22

Control 0.94 + 0.03 0.84 0.95 0.96 + 0.00 0.94 + 0.02

212 Table IV-7. Diversity as measured by Fisher index ( ) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited / Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 9.50 + 2.34 21.31 + 3.21 22.46 + 3.54 20.27 + 4.30 25.01 + 7.26

Control 29.28 + 7.52 36.04 46.60 44.67 + 6.56 41.03 + 6.26

Table IV-8. Diversity as measured by Taxomonic Diversity index ( ) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited / Preharvest Post 1 Post 2 Post 3 Post 4/5 Control

Baited 14.22 + 7.68 11.29 + 4.09 11.85 + 3.28 1 6.47 + 5.38 36.04 + 15.66

Control 67.04 + 2.21 52.75 69.7 1 67.20 + 1.57 62.02 + 2.74

Table IV-9. Diversity as measured by Taxonomic Distinctness index ( *) from baited and unbaited pheromone trapping sites in Douglas-fir habitat, preharvest through 4/5th season post harvest conditions ( = 0.05). Pheromone traps baited with components MCOL, seudenol, and frontalin known to aggregate Douglas-fir beetles. Post 1 and post 2 unbaited data represent a single trapping site. Baited/ p reharvest Post 1 Post2 Post 3 Post 4/5 Control

Baited 63.01 + 8.65 65.73 + 1.98 64.33 + 6.89 71.51 + 0.67 71.49 + 0.96

Control 71.24 + 0.95 63.00 73.17 70.03 + 1.41 66.16 + 2.13

213