UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

The biodiversity of flying Coloptera associated with integrated past management of the Douglas-fir beetle… Carson, Susanna Lynn 2002

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-ubc_2002-0350.pdf [ 10.63MB ]
Metadata
JSON: 831-1.0090382.json
JSON-LD: 831-1.0090382-ld.json
RDF/XML (Pretty): 831-1.0090382-rdf.xml
RDF/JSON: 831-1.0090382-rdf.json
Turtle: 831-1.0090382-turtle.txt
N-Triples: 831-1.0090382-rdf-ntriples.txt
Original Record: 831-1.0090382-source.json
Full Text
831-1.0090382-fulltext.txt
Citation
831-1.0090382.ris

Full Text

THE BIODIVERSITY OF FLYING COLEOPTERA ASSOCIATED WITH INTEGRATED PEST MANAGEMENT OF THE DOUGLAS-FIR BEETLE (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 p resent ing this thesis in part ial fu l f i lment of t h e requ i rements f o r an advanced degree at the Universi ty of Brit ish C o l u m b i a , I agree that t h e Library shall make it f reely available fo r re ference and s tudy. 1 fu r ther agree that permiss ion fo r extens ive c o p y i n g o f this thesis f o r scholar ly pu rposes may b e g ran ted by the head of my d e p a r t m e n t or by his o r her representat ives. It is u n d e r s t o o d that c o p y i n g o r pub l i ca t i on o f this thesis fo r f inancial gain shall n o t be a l l o w e d w i t h o u t m y w r i t t e n permiss ion . D e p a r t m e n t The Univers i ty o f Brit ish C o l u m b i a 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 species management on local insect 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 4/5 t h 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' 1 0), 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 Sampl ing Techn iques 8 Trap design 8 Sampl ing Design 10 Trap placement 10 Replication 12 Site Condi t ions 14 Topography/Climate and weather 14 CHAPTER II - Impact of Aggregation Pheromones on Old Growth Associated Flying Beetle Diversity 17 Introduct ion 17 Methods 21 Resul ts 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 Introduct ion 65 Methods 69 Species distribution analysis 74 Diversity analysis 75 Species abundance - baited vs. control 76 Species trends 77 Resul ts 77 Diversity analysis / Overview 100 Spec ies Number (S) 101 Marqa le f s (d) 103 Pielou's (J') 104 Bril louin 104 Shannon-Wiener (H'ig) 105 S impson 's M-X) 106 Fisher's (a) 107 Taxonomic Diversity (5) 107 Taxonomic Dist inctness ( 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 Harvest ing summary 125 Pheromone Effect 128 Natural var iat ion 136 Trapping eff iciency 137 Systemat ic bias 139 iii Semiochemica l variat ion 140 Summary 140 CHAPTER IV - Pheromones and Integrated Pest Management 142 Introduct ion 142 Pheromones , Insect Ecology, and Communi ty Deve lopment 146 M a n a g e m e n t Impact 148 Containment and mop-up 148 Monitoring 151 Pheromone bias as context 152 Sources of Var iat ion and Exper imental Limitat ions 155 Species variation 155 Habitat variation 157 Study limitations 158 Pheromone composition 159 Site selection and replication 160 Summary 162 BIBIOGRAPHY 166 Append ix I Taxonomic suppor t 183 Append ix II Statist ical protocols for Douglas-f i r beetle abundance 185 Append ix III Species data by t rend 187 Append ix IV Tabular results, Chapter 3 diversity indices 210 iv List of Tables* Tab le 1.1 Site repl icat ion (preharvest condi t ions) 14 Table 2 . 1 List of non-target species known to aggregate to Douglas-f i r beet le phe romone componen ts 20 Table 2.2 Site list, t rapping year, and biogeocl imat ic classi f icat ion for preharvest phe romone baited and unbai ted sites 23 Table 2 .3 S u m m a r y results of mean abundance/s i te and total abundance of f ly ing beetle species under preharvest condi t ions 30 Tab le 2.M Twenty most abundant f lying beetle species (preharvest) 38 Table 2 . 5 Richness, evenness and dominance measures of f lying beet le diversi ty (preharvest) 41 Table 2M Spec ies of f lying Co leoptera statistically more or less abundan t be twee pheromone baited and control sites (preharvest) 42 Tab le 3.1 Site list, t rapping year, and biogeocl imat ic classif icat ion for preharvest and postharvest pheromone baited and unbai ted sites 69 Table 3.2 Site repl icat ion of baited and control data across t reatment years 74 Table 3.3 Site list for 5-repl icate diversity analysis, including harvest stage, t rapping year, and biogeocl imat ic classif icat ion 76 Table 3.H S u m m a r y results of total abundance and species number of f lying beet les (preharvest and postharvest) 79 Table 3.5 S u m m a r y results of mean abundance/s i te and total abundance of f lying beet le spec ies (preharvest and postharvest) 80 Table 3.6 Ten most abundant beetle species listed by rank f rom phe romone baited and control sites 99 Table 3.7 Flying Coleoptera species statistically more or less abundant be tween pheromone baited and control sites (preharvest and postharvest ) 109 T a b l e tit les have been abbrev ia ted/paraphrased for length. v List of Figures* Figure 1.1 Distr ibution of Douglas-f i r in North Amer ica 1 Figure 1.2 Northern Distr ibution of mature Douglas-f i r 2 Figure 1.3 Site Map 13 Figure 2 . 1 Site Map - preharvest condi t ions 22 Figure 2.2 D iagrammat ic representat ion of sampl ing design 25 Figure 2.3 V e n n d iagram of species distr ibution (preharvest) 36 Figure 2M Whi t taker plot of species abundance (preharvest) 37 Figure 2.5 Whi t taker plot of 20 most abundant species (preharvest) 37 Figure 2 .6 Resul ts of f lying beetle diversity measures (preharvest) 40 Figure 3.1 Mean abundance/s i te of f lying beet les in pheromone bai ted and control si tes (preharvest and postharvest) 96 Figure 3.2 Whi t taker plot of species abundance (preharvest and postharvest ) 98 Figure 3.3 Resul ts of f lying beetle diversity measures in phe romone baited and control si tes (preharvest and postharvest) 102 Figure 3. 1/ Observed t rends in f lying Coleoptera t rapped in phe romone bai ted t raps under preharvest and postharvest condi t ions 111 Figure 3.5 Observed t rends in f lying Coleoptera t rapped in unbai ted contro l t raps under preharvest and postharvest condi t ions 111 Figure 3.G Success ive phases in the invertebrate communi ty exploi t ing dead w o o d 115 Figure 3.7 Phases of ecosys tem deve lopment after clear-cutt ing of a second growth northern hardwood forest 126 *Figure tit les have been abbrev ia ted/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 insects, 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 taxonomy 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. Figure 1.1. Distr ibut ion of Douglas-fir in North Amer i ca ( • ) adapted f rom Hermann and Lavender (2001). 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 2001). In the Fort St James Forest 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). Figure 1.2. Distr ibut ion of the northern limit of mature, leading (>f 8 ) Douglas-f i r (Pseudotsugae menziesii) in the Fort St James Forest District, Brit ish Co lumbia , a long with current Interior Douglas-f i r ( IDF) biogeocl imat ic ecozone designat ion ( • ) . Inset shows southwest boundary of the Fort St J a m e s Forest District, major lakes, and leading Douglas-fir (>f 8 , >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 IDF Biogeoclimatic ecozone 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 Curculionidae 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 pine, spruce, 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 genus 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 arthropods 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-hickory 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, 1 s t season post-harvest, 2 n d season post-harvest, 3 r d season post-harvest and 4 / 5 t h 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 animal damage. 12 C P 32/80 (1) C P 120/2 (3) C P 46/222 (3) K u z Che (1) C P 123 (3) C P 118/1 (3) C P 115 (6) Hobson Is. (5) TachieHil l (5) Germajisen-Pinchi (10) P inch iHi l l (2 ) Tachie-Pinchi (5) R N W (2) R N E (2) A P 1 - 4 , A P C , C P 1 8 (10) Whitefish(5) Siesmic (1) Figure 1.3. Locat ion of 17 study areas, containing 67 sites, initiated between 1994 and 1997 in the Fort St J a m e s Forest District, British Co lumbia . The number of si tes 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. Tab le 1.1. Site repl icat ion of baited and control data across t reatment years for the Fort St J a m e s pheromone study. * Indicates sites in wh ich diversity ca lcu lat ions are inc luded for genera l compar ison only. T rea tmen t year Preharvest Post 1 Post 2 Post 3 Post 4/5 Bai ted or control bait control bait control bait control bait control bait control # of site repl icates 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 S p e c i e s Beetle attacked Douglas-fir 1 Douglas-fir volatile +EtOH 2 M C O L 3 Fn 4 S e u d -enol 5 References 1-5 Dendroctonus pseudotsugae Hopkins (Scolytidae) • # • • • u , J , 4 Wood 1982 4 Dickens ef al. 1985, Lindgren 1992. 5 Pitman etal. 1975, Ross and Daterman 1995. Dendroctonus brevicomis Lec. (Scolytidae) » 4 Wood 1982. Dendroctonus frontalis Zimm. (Scolytidae) • Pine 4 Wood 1982, Payne et al. 1978, Payne etal. 1988. Dendroctonus ponderosae Hopkins (Scolytidae) • Pine J Borden etal. 1990 (contrary to Libbey et al. 1985) Dendroctonus rufipennis Kirby (Scolytidae) • Spruce • Spruce • Spruce J Borden etal. 1996 4 Dyer 1973, Lindgren 1992. 5 Furniss etal. 1976. 3 , 4 , 5 Setter and Borden 1999. Dendroctonus terebrans (Olivier) (Scolytidae) # Pine 4 Payne etal. 1987, Delorme and Payne 1990. Hylastes nigrinus Mann. (Scolytidae) • • 'Rudinsky & Zethner-Moller 1967. 2Rudinsky 1966a. Hylates ruber Swaine (Scolytidae) • ^Rudinsky 1966a. Trypodendron Lineatum Oliv. (Scolytidae) • • z Rudinsky 1966a. 4 Lindgren 1992, Setter and Borden 1992. Dryocetes autographus Sw. (Scolytidae) m ^Rudinsky 1966a. Gnathotrichus • ^Rudinsky 1966a. 20 retusus Lec. (Scolytidae) 9 Gnathotrichus sulcatus Lec. (Scolytidae) • ^Rudinsky 1966a. Pseudohylesinus grandis Sw. (Scolytidae) • 'Rudinsky 1966a. Pseudohylesinus nebulosus LeC. (Scolytidae) z Furniss et al. 1974, Rudinsky 1966a. Thanasimus dubius LeC. (Cleridae) • Pine 4 Wood 1982, Dixon & Payne 1980. Thanasimus undatulus Say. (Cleridae) • ' Furniss etal. 1974. 4 Wood 1982, Lindgren 1992. Enoclerus sphegeus Fab. (Cleridae) • 1 Furniss ef al. 1974. Abraeus sp (Histeridae) # Pine 4 Dixon & Payne 1980. Cylistix attenuata (Histeridae) • Pine 4 Dixon & Payne 1980. Platypus flavicomis Herbst (Platypodidae) • Pine * Dixon & Payne 1980. 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 18 (1) Whitefish (2) Siesmic (1) Germansen-Pinchi (2) Tachie-Pinchi (2) Kuz Che (1) CP 118/1 (2) CP 115(1) CP 120/2 (2) F i g u r e 2 . 1 . Locat ion of preharvest study areas, initiated between 1994 and 1997 in the Fort St J a m e s Forest District, Brit ish Columbia. Dots indicate all majori ty ( F 8 or greater) , mature to over mature Douglas-f i r in the Fort St James Forest District. Sol id shading indicates location of major water bodies. Left boundary line indicates western limit of forest district. The number of si tes per s tudy 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 3cm 2 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 (>Fd 8), 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, t rapping year, and biogeocl imat ic classi f icat ion for p reharvest p h e r o m o n e baited and unbaited sites in the Fort St J a m e s Forest District, Brit ish Co lumb ia Site Number Site Name/Locat ion Sampl ing Year Biogeocl imat ic sub z o n e 1 Tachi -Pinchi 1994 S B S d w 3 1c Tachi -Pinchi 1997 S B S d w 3 2 Germansen-P inch i 9 km 1994 S B S m k 2c Germansen-P inch i 8 km 1997 S B S m k 3 T F L CP 115 1996 S B S w k 3 4 T F L CP 120 1997 S B S d w 3 4c T F L CP 120 1997 S B S d w 3 5 T F L CP 118 1996 S B S w k 3 5c T F L CP 118 1997 S B S w k 3 6 Kuz C h e 1996 S B S w k 3 7 Whi tef ish D 1997 S B S w k 3 7c Whi tef ish D 1997 S B S w k 3 8c Apol lo CP 18 1997 S B S w k 0 1 / 0 4 9c Seismic Trail 1997 S B S d k 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 o f 1 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 o f 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 o f 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 2 6 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 <x has low sensitivity to sample size. Assuming the selected range of species richness indices would be sufficient to account for sampling intensity, additional indices were then added to emphasize dominance (1-Simpsons), and evenness (Pielou's J'). The study also took the opportunity to include two recently described indices; Taxonomic diversity (6), and Taxonomic distinctness (6*). Both indices integrate a measure of "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 (Ciidae); 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 Bradycellus nigrinus (Dejean) 0.1 0.0 1 Calathus 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 Cephaloon 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 3 0 (leConteJ Tetropium 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 Cucujidae Cucujus claviceps Mannerheim 4 1.9 53 Dendrophagus cygnaei Mannerheim 1.7 0.3 19 Pediacus 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 Dermestidae 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 Erotylidae 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 Leiodidae 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 Melandryidae 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 Mycetophagidae 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 Salpingidae Rhinosimus vindiaeneus Randall 2.1 o.i 22 Scaphidiidae Scaphisoma castaneum Motschulsky 0.1 0.0 1 Scarabaeidae 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 Scirtidae 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 Scraptiidae Sphindidae Staphylinidae 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) 1 2 7 4 1 3 . 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 Anaspis sp. 2 5 . 6 1.4 266 Hallomenus sp. 0.1 0 . 3 3 Orchesia (nr.) castanea 0.1 0.1 2 Odontosphindus clavicornis Casey 0 0.1 1 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)) 2 0 2 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 Tenebrionidae Tetratomidae Throscidae Trogossitidae (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 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 Tetratoma concolor LeConte 1.9 1.4 29 Pactopus 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 3 5 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: 3 1 % (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. V e n n d iagram showing the spec ies distr ibution of f ly ing Co leop te ra found in funnel t raps baited for Douglas-f i r beet le in the Fort St J a m e s Forest District, Brit ish Co lumbia . Habitat is Mature/Old growth Douglas-f i r (Fd 8, >115 yrs). Data exc ludes identif ied necrophagous spec ies or spec ies <1 m m . 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 — « - N n t i f l ( D N f l O ( i ) O T - N ( i ) ^ i n < O N e o o ) o T - T - T - T - i - r - T - T - r - r - C N J Species Rank (1-20) 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 composit ion 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 (Fd 8, >100 yrs). Listings do not include identified necrophagous species. Rank #/si te Preharvest Baited Twenty Most Abundant Species #/site Preharvest Control t 7890Dendroctonus pseudotsugae Hopkins 2 ^QAATrypodendron Lineatum (Olivier) 3 285Eusphalerum spp (mostly pothos (Mannerheim)) H 163 Thanasimus undatulus (Say) 5 31Anaspis sp .(? 17 Hylastes nigrinus (Mannerheim) 7 15Scierus annectans LeConte <J BCorticaria n sp S IXylechinus montanus Blackman 10 SQuedius velox Smetana 11 6Pseudohylesinus nebulosus LeConte 1Z 5Hylastes ruber Swaine 1 3 5Pelecomalium testaceum (Mannerheim) 1H 5Quedius plagiatus Mannerheim 1 5" ASpondylis upiformis Mannerheim I (c AAgathidium depressum Fall /obtusum Hatch II ACtenicera pudica (W.J. Brown)+ propola columbiana (Leconte) APolygraphus rufipennis (Kirby) 1 °t AEnicmus tenuicornis LeConte 20 3 Tetratoma concolor LeConte \7>Trypodendron Lineatum (Olivier) 7Hylastes ruder Swaine (Mannerheim)) 7Eusphalerum spp (mostly pothos Mannerheim)) 6Dendroctonus pseudotsugae Hopkins GHylastes nigrinus 5Ctenicera pudica (W.J. Brown) +propola columbiana (Leconte) 5Thymalus marginicollis Chevrolat 5Quedius velox Smetana APseudohylesinus nebulosus LeConte 4Scierus annectans LeConte ZBolitopunctus muricatulus (Hatch) ZTrichochrous albertensis Blaisdell ZSpondylis upiformis Mannerheim ZEnicmus tenuicornis LeConte ZDrasterius debilis LeConte ZAgathidium depressum/obtusum Fall/Hatch ZAmpedus brevis Vandyke ZAthous rufiventris rufiventris (Eschscholtz) 2Ctenicera nebraskensis (Bland) 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 (<x) 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 55 s. • • 1 Value, o 10 - WL * i • Calculated O) o 1 -0 . 1 . -! i i c 0 01 s I I I I I N ^ Jyi *2> JL Diversity Measure/Index Figure 2.6. Measures of f lying beet le diversity f rom phe romone bai ted ( • ) and unbai ted/contro l (a) si tes in interior Douglas-f i r (Fort St J a m e s Forest District, Brit ish Columbia) . Beet les were t rapped in L indgren funnel t raps. P h e r o m o n e bai ted t raps were baited with a ternary lure of M C O L , seudeno l and frontal in. Si tes w e r e Mature/Old growth Interior Douglas-f i r (Fd 8 , >100 yrs). Indices do not include identif ied necrophagous species or spec ies < 1 m m . 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 (Richness) 58.3 + 12.5 56.0 + 8.02 0.30 0.769 Margalef, d (Richness) 6.29 + 1.29 10.9 + 1.35 -4.87 0.000 Shannon, H' (Richness) 0.197 + 0.0861 1.51 + 0.0984 -25.97 0.000 Brillouin (Richness) 0.444 + 0.198 3.04 + 0.194 -18.37 0.000 Fisher, a (Richness) 8.39 + 1.94 32.7 + 6.41 -7.12 0.000 Pielou, J' (Evenness) 0.11 + 0.0436 0.87 + 0.0372 -19.75 0.000 1-Simpson, 1-D (Dominance) 0.194 + 0.126 0.952 + 0.0208 -11.64 0.000 Taxonomic diversity, 8 15.8 + 8.65 67.99 + 1.93 -11.54 0.000 Taxonomic distinctness, 5* 61.8 + 6.22 71.5 + 0.772 -3.03 0.023 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. Spec ies of f lying Coleoptera statistically more or less abundan t be tween phe romone bai ted and unbai ted control si tes in mature to overmature interior Douglas-f i r under at tack by the Douglas-f i r beetle. P h e r o m o n e bai ted t raps w e r e bai ted wi th a Frontal in, M C O L , Seudeno l lure known to aggregate Douglas-f i r beet les (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 Draster ius debi l is 11.5 14 S C O L Y T I D A E E L A T E R D I A E Thanasimus undatulus 1 14 eanthophlax mirificus 9.5 • 14 C L E R I D A E E R A M B Y C I D A E Anaspis sp. 14 14 STAPHYLINIDAE Polygraphus rufipennis 13.5 14 S C O L Y T I D A E Rhizophagus dimidiatus 13 14 R H I Z O P H A G I D A E Rhinosimus viridiaeneus 13 14 S A L P I N G I D A E 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 communi ty 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 communi ty responding to a pheromone lure consists of all species exhibit ing 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. Whi le it may be more ecologically appropriate to define such a communi ty 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 conifers (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 5 2 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 f rontal in, 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. 6 4 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 ground beetle 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). Tab le 3 . 1 . Site list wi th harvest stage, t rapping year, and b iogeocl imat ic c lassi f icat ion for pheromone-ba i ted and unbaited sites in the Fort St J a m e s Forest District, Brit ish Co lumbia . Subzone defini t ions can be found in Meid inger and Pojoar (1991) . Site N a m e Biogeo-cl imatic Subzone Preharvest P o s t - 1 P o s t - 2 P o s t - 3 Post -4/5 Bait Control Bait Control Bait Control Bait Control Bait Control 1 T a c h i e H i l l S B S d w 3 94 95 96 97 97 2 Tachi -P inch i S B S d w 3 94 97 95 96 97 3 G P 8 & 9 k m S B S m k 94 97 97 4 G P 13 k m S S B S m k 94 95 96 97 97 5 G P 13 k m N S B S m k 96 97 6 A P C / C P 1 8 SBSdk 97 96 97 7 Hobson Is. S B S d w 3 94 95 96 97 97 8 A P 1 - 4 S B S d w 3 95 96 96 97 97 97 97 9 W D S B S d w 97 97 95 97 97 10 S iesmic SBSdk 97 11 R N E SBSk3 97 97 12 R N W S B S k 3 96 97 13 Pinchi Hill S B S d 0 6 97 97 14 C P 123 96 97 97 15 Kuz C h e S B S w k 3 96 16 CP 32-80 96 17 CP 46-222 S B S w k 3 96 97 97 18 C P 115-1 S B S w k 3 96 96 97 97 19 115 (Randy) S B S w k 3 97 97 20 C P 118 S B S w k 3 96 97 97 21 C P 1 2 0 S B S d w 3 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 3cm 2 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 ( Fd 8 ) 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 i 1) 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 o f 1 Fn was 0.3 mg/day from capillary tubes of 1.0 mm diameter. The MCOL-seudenol blend was achieved by dispensing p u r e 1 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, 1 s t 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 repl icat ion of baited and control da ta across t rea tment years for the Fort St J a m e s pheromone study. * Indicates sites in wh ich diversity ca lcu lat ions are included for genera l compar ison only. T rea tmen t Preharvest Post 1 Post 2 Post 3 Post 4/5 year Bai ted or control bait control bait control bait control bait control bait control # of site repl icates 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, 1 s t , 2 n d , 3 r d , and 4 t h /5 t h season postharvest conditions and five control sites for each of preharvest, 3 r d , and 4 t h /5 t h season postharvest conditions. Single control replicates for 1 s t , and 2 n d 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'i 0), 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 inc ludes harvest s tage, t rapping year, and biogeocl imat ic classif icat ion (Fort St J a m e s Forest District, B. C ) . Site N a m e Biogeo-cl imatic Subzone Preharvest Post -1 Post -2 Post -3 Post -4/5 Bait Control Bait Control Bait Control Bait Control Bait Control Tach ie HII S B S d w 3 94 95 97 97 Tachi -P inch i S B S d w 3 94 97 95 96 G P 8 & 9 k m S B S m k 94 97 97 G P 1 3 k m S S B S m k 94 95 96 97 97 Hobson Is. S B S d w 3 94 95 97 97 A P 1 - 4 S B S d w 3 97 97 97 97 W D S B S d w 3 97 97 97 97 R N E SBSk3 97 97 Pinchi Hill S B S d 0 6 97 97 CP 123 96 97 97 CP 46-222 S B S w k 3 97 CP 115-1 S B S w k 3 96 97 C P 1 1 8 S B S w k 3 96 97 C P 1 2 0 S B S d w 3 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 7 6 subject to non-parametric statistical analysis by Wilcoxon Rank-sum test. Harvest years with only one control replicate ( 1 s t and 2 n d season postharvest) were omitted from this analysis. Species trends Using baited and unbaited data sets, mean species abundance was recorded from preharvest, and 1 s t , 2 n d , 3 r d and 4/5 t h 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 Tab le 3.4. Summary results of abundance and spec ies number of f ly ing beet les occurr ing in bai ted and unbaited Lindgren funnel t raps in mature and recent ly harvested Douglas-f i r habitat (1,2,3,4/5 years postharvest) (Fort St J a m e s Forest District, Brit ish Columbia) . Baited funnels t raps conta ined phe romone lures for the Douglas-f i r beet le {Dendroctonus pseudotsugae) consist ing of M C O L , seudeno l , & frontal in. T rea tmen t # Site repl icates (3 trap/site) Total # of species Total # of beet les Total # Douglas-f i r beet les # of non-target spec ies 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 Al l si tes 65 625 484,319 380,903 103,415 7 9 Table 3.5. S u m m a r y results of mean abundance per site and total abundance of f lyng beet les occurr ing in baited and unbai ted Lindgren funnel t raps in Mature and recent ly harvested Douglas-f i r habitat (1,2,3,4/5 years postharvest ) (Fort St J a m e s Forest District, Brit ish Columbia) . Baited funnels traps conta ined pheromone lures for the Douglas-f i r beetle (Dendroctonus pseudotsugae) consist ing of M C O L , Seudeno l , & Frontal in. Species with a mean abundance/s i te >100 have been rounded to the nearest who le number . Family Genus species in CD 2 ro .c CD CL B B B -5? 'co c "c "c ro ro ro CD CD CD E, E, E, TJ T 3 "O B B ' ro ' ro ' ro .O n - 0 CM m to to to o o o CL CL CL 0.0 0.1 0.0 0.0 0.0 0.0 B •£ (/) 03 ro t CD ^-E p ro to o 0-CD CL CD E o CL CD E o o CM O CL CD E o o CO o CL B ro CD E. o T 3 CD C Q . O CL o ro ID ?f to 3 O o O. r-0 0 1 0.1 1 Alleculidae Anobiidae Anthribidae Bostrichidae Buprestidae Byrrhidae Byturidae Isomera (nr) comstoki Papp sp. # 1 Caenocara scymnoides LeConte Desmatogaster subconnata (Fall) Dorcatoma (prob) americana # 87 Ernobius gentilis Fall Ernobius nigrans Fall Hadrobregmus americanus (Fall) Hadrobregmus quadrulus (LeConte) Hemicoelus carinatus (Say) Microbregma e. emarginatum (Duftschmid) Ptilinus lobatus Casey /basalis Leconte Utobium elegans (Horn) Xyletinus rotundicollis R.E. White Allandrus populi Pierce Tropideres fasciatus (Olivier) Stephanopachys substriatus (Paykull) Anthaxia inornata (Randall) Buperstis langi Mannerheim Buprestis lyrata Casey Buprestis nuttalli Kirby Chrysobothris carinipennis LeConte Dicerca tenebrica (Kirby) Dicerca tenebrosa (Kirby) Melanophila drummondi (Kirby) Curimopsis sp. Bhyrrhus sp. Cytilus sp. (alternatus) (Say) Byturus unicolor Say .0 0 .0 0.0 0.0 0.0 0.0  .0 0.0 0.0 0.0 0.0 0.0 0  0.2 0 .0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0 .2 0.0 0.0 1.0 0.0 0.0 0.4 0.0 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 6 0.0 0.4 0.0 0.0 0 .2 0.0 1.0 0.0 0.0 0.6 9 0.0 0.1 0.0 0 .2 0.0 0.0 0.0 0.0 0.0 0.0 3 0.0 0.3 0.5 1.1 0.0 0.0 0.0 0.0 0 .2 1.4 2 8 0.0 0.0 0 .2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 0.0 0.1 0.1 0 .2 0 .2 0.1 0.0 0.0 0 .2 0.0 7 0.6 2 .3 2 .4 0.7 0.0 0.7 1.0 6.0 0.4 0 .2 0.0 0 .0 0.0 0.0 0 .2 0.0 0.0 0.0 0.0 0.0 0.1 0 .0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0 .0 0 .2 0 .2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.0 7 9 1 3 4 3 0.0 0 .2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 0 .2 1.7 0.5 0.3 0.0 0.1 0.0 1.0 0 .2 0.0 3 2 0.0 0.0 0.0 0 .2 0.3 0.0 0.0 0.0 0.0 0.4 6 0.0 6.5 4 . 5 4 . 5 6 .2 0.0 0.0 11 .0 8 .4 7.2 2 9 1 0.0 0 .2 0.7 0.3 1.0 0.0 0.0 0.0 0.0 0.6 2 2 0.0 0.0 0.3 0.4 0.0 0.0 0.0 0.0 0.6 0.0 10 0.0 2 .2 9 .4 9 .6 7.5 0.0 1.0 3.0 12 .0 8 .8 3 7 6 0.0 3.3 3.0 0.9 1.0 0.0 5 .0 0.0 1.6 0.4 9 9 0.0 0.5 0.4 0 .2 0.0 0.0 1.0 0.0 0.4 0.0 14 0.1 0.0 0.1 0.0 0 .2 0 .0 0.0 0.0 0.0 0 .2 4 0.0 0.1 0.1 0.0 0.0 0.1 0.0 0.0 0.0 0.0 3 0.0 0.3 0.5 0 .2 0.7 0.0 1.0 0.0 0.0 0.2 17 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 Elaphrus 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 Lebia 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 Cerylonidae Chrysomelidae Ciidae Clambidae Cleridae Coccinellidae 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 Cerylon castaneum Say 0.6 0.8 1.3 0.9 0.7 0.9 2.0 2.0 0.8 0.4 58 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 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 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 Adalia bipunctata 0.0 0.0 0.2 0.8 0.3 0.0 0.0 0.0 0.0 0.2 13 83 Colydiidae Corylophidae Cucujidae Curculionidae (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 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 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 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 Pediacus depressus (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 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 8 4 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 Derodontidae 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 Sericus 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 Epiphanis cornutus 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 3 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 Hydraenidae 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 Helophorus 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 o:o (Linneaus) 0.0 0.0 0.3 0.0 0.0 0.0 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 Typhaea stercorea (Linneaus) 0.0 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2 Nemonychidae Cimberis (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 Ptinidae 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 Sphaeriestes 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) Cyphon concinnus 0.1 0.3 1.1 1.5 0.7 0.0 0.0 0.0 2.4 1.0 52 (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 Sphaerites 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 Aleocharinae (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 Stenotrachelidae 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) Corticeus praetermissus 0.0 1.5 0.3 0.0 0.0 0.3 0.0 0.0 0.2 0.0 23 (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 Trixagus 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) Ostoma ferrugina 0.4 1.3 2.0 4.7 4.3 0.4 0.0 5.0 1.0 1.8 135 (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 "55 o o a> CO * c O 1000 100 Preharvest Post 1 Post 2 Post 3 Harvest Stage Post4/5 Figure 3 . 1 . A b u n d a n c e of f ly ing beet les in pheromone-ba i ted ( • ) and unba i ted* ( • ) L indgren funnel t raps t rapped in p reharves t th rough 475 t h season post harvest condi t ions. Habi tat condi t ions were s tand ing , or recently harves ted , Douglas- f i r beet le a t tacked, mature Inter ior Douglas- f i r ( F d 8 , >100 yrs), Fort St J a m e s Forest District, Brit ish Co lumbia . * Post 1 and post 2 unbai ted data represent a single t rapp ing site, and are inc luded for genera l compar i son 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 1 s t 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 415 th postharvest year). As Scolytids decreased, members of the family Elateridae increased, comprising at least half of the top ten species in 4/5 t h season postharvest conditions (5/10 species & 7/10 species: baited & control sites respectively). 97 10000 1000 100 10 • Preharvest Baited Preharvest Control Species Rank (1-208) a. Preharvest 100000 10000 1000 100 Post-1 Baited Post-1 Control Species Rank (1-416) b. First season p o s t h a r v e s t ' 100000 10000 1000 100 10 1 0.1 0.01 • Post-2 Control —* • Species Rank (1-413) c. Second season postharvest * -Post-3 Baited -Post -3 Control Species Rank (1-350) d. Third season postharvest 10000 1000 100 10 1 i—Post-4/5 Control Species Rank (1-262) e. Fourth/f ifth season postharvest Figure 3.2. Species abundance of f lying Coleoptera in a) preharvest habitat, b) 1 s t season postharvest habitat, c) 2 n d season postharvest habitat, d) 3 r d 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 phe romone-baited and unbai ted/control sites. Beet les were t rapped in L indgren funnel t raps in mature Interior Douglas-f i r ( F d 8 , >100 yrs) in the Fort St J a m e s Forest District, Brit ish Co lumbia . * Control data result ing f rom a single site inc luded for general compar ison only. 98 CD q a. m 3 R s-> 3 § S'l 3 m c m Q) o — Cr ^ 00 cn CD 3 ' 03 o r o o oo § rO CO CIS 3. 3 C Co tn 9J Co 3 2-=• d 0 3 M o ^_ 3 CD 3 5 c 2 tn C Co O c CD Q. c Ji w c J o a , S O -Qi - cp CQ . 1 § tn m o *s o n> q a. 5 co fn 3. ^ f n m z£ CD m § S g fn O ^ O 0) q a. 5 " > 3 m C ^- o-CD 9 "° O o c y g" & 3 18 I 9^  + CD m D ) 5 -— a O oT ro » 2 5 o a» o o n CD <Q 3 §1 g m c: ro f 3 1 . : <s' 2 I 2 <D ro m Z 0) > «a m 03 s i co S5. P CD 3 § 5 3 « 3 n-s-g g co O 3 m Si =: ^ 3 S 03 3 ! o c > co m It | m i l 5 § 5T 5T Cr 3 3. 3 03 O s "S' 9- 3" i i ro 2 2 (5" -£ 3 o >r o o 2. 3- 5 co Co E r 2 i» ; "9' 3 2 <D ' to m z a> » ST 5 2 a 5. > CQ " nT m (Q If l ! c § _ c CD Co 2 . 3 ' "co O Q) q a H CD 5 M > 3 Bg-CD z a> tj 5. > <Q m 0 ) 55" fjT c? O c 3 cV 10 CD CD -1§ 0 ) s l l g CO' 2 > <Q "S > 3. sr ; 3 co • 2 " D l CD tn cn • s> to 3 2-a s ST " 3 S o o c — n 3 Ifl r & C J m (D* 0) - ° Co 3' c 5 a > tQ m Q> a . to c -•m 2. O ,„ 2 o > .-3 5 3 c CO co 3 O s- 3 o S n> -< c 3-3 co o > O' c CO ^ 3 . 1 O (Q "S O 3. 0) q C " 3 « S > m •m § O § 1 ! | 8'8' g 5T S ~m CD * i co S2> m ? oT m 2. 51-1 m 2 2 § g & > «» m C J 3 i. , | i m a g > w m ' -6 o o 13 d co o D C S ' > Co ™ CD CO 3' c Co cn 3 (Q CD 03 CD 3 , co Co 'co\3l o *s O Qj q -1 CD C • o-CD III 5 i t • &r a £ c o mtQ 3 03 c CD Co 5t § S § 8' g &s m 3 m" 3 O a § 8' g S-S m 3 ^— Co f n 5 O a § 8' g | S m 3 •—' co ' « 2 .1 O 1Q O 3. 03 r 3 S P cf Co Co Ol o> • a> 3 03 tn O ? § 3 m Q. 03 5 » -a c > &• • 03 . Co CO ^ o "s O 03 q --) CD 5 S 3 -2-lQ' 3. 3 C O Ol o 2 "s o d e q 8<a > Co m CO sit • sr a £ c o m CQ 3 03 c CD Co sp . m CO o 3 £" q gca = to & m co 8 ^ 2 T J vu Co CD" CD ~ c 3" S-co g . to "S. CD Co 5 a z a> > (Q m 03 2 J Q. 03 03 > 3 co-co to o «• o > CD m CD 2 3 B- m 3 3 3 8 2 P-3 " • Co ' ~ -. 03 y S'-Q > 3 3 03 ' "O o oT ro o ro 5! "CS S - D X T3 3-•< O 03 £ 3 CD" 2 o 3 > T 3 co i n J -< O 03 i l l g ^ i •co ^ " ^. O 03 q a -1 CD 5 co BCQ 3 3' C Co o £ 3 •SCQ' 3. 3 C CO O 03 ; 3 '<3' 3' c Co m g CD g CO m S 1 1 2 -if g 5T co 3 Co o CD , 3 3 & 3 g | « 3 5 3 ro 'tn ^ ^ 3 1 & O 03 CD o to > 3 B & CD O 3 ' ^ CD H ^0 03 a a- o 0 1 CO 00 "tn ^Z. 'tn o "S O 03 q a q CD 3 I 3 " 2 § | - 0 T 3 d co' O Co «• 2.3: tr> CQ "s o 3. 5T q g a 3 « a > m 'tn <"> ^ . O 03 q a H CD 6 co CQ' 3 3' C CO CO > 3 CD ? -Q' 3 5 2 O' 2 3 CD D CD 1 > 3 i m CD ~ C3- i 3.-o 03 ro (D ->• ro CA3 O ; S.8 8"g Q cr 3 a. 5 £-5.2 5. s a H 5 2 ^ 9- o 5 2. a! o" CD CD sj. 2 2^  + CD i | S t g | | s co' Co' C m 03 O a o ST 03 ov 3 8 ? 0) i> c 9. o §111 Q. _ 71 03 3 H I co 3 c Co o' m 2 H CD g §'•§ 2 m 03 P O ST ro Ol I 9 l l O' C 3 8 2 OT 03 ro co m ctj 5t m CD 7> 3 5 9-> 2 m 3 ^ co 2 3 CD o 3 > m 03 ?c3 5 ? S ? 0 5 Cr CD 5 H 3 3 H m o> s- m 2 % 8 2 3. Co a c ~? 5- a § I? 8 - § " § s m cu o O S" ai co co • I o CD , c ^ I O Co > c m CQ 03 1% 1 0 3 fo §^ rf| o n, -g ro: co 0 1 CD , C d & O tn > C m CQ OS fn 31 ^ O 03 ^ §• 3 00 C3 f" 31 ^ 8 l l 5 = S Ol Ji ' 2 "0 CD CO C3- CD £ C O & CO O C 3-I | | 5 co a > c o rncQ 3 03 c CD Co -J o m 5 a Z 03 > "9 m 03 "~' sr Ji Ol ill; s to a > c o m c Q 3 03 C CD Co fn § Q! § I 8" g or 3 HI 5 It' 5 co a > c o m c Q 3 03 C CD Co § 9 § 2 o' f5 o 2. ST 03 71 7? 5 S| CO CD CD => A CD' ^— Co S 2 CO CD CD 3 * 2 CO CD CD 3 •S.CO ct> Q A » ' Co CO CD CD 3 3f d CD <S CO CO CD g-' <» CD 3 cf Q A CD ^— co CO CD 5-' *> CD 3 6 CD CO CO CD 3-' a CD 3 f?.co 9>. ID" CD Q A K' Co CO CD S-' *> CD 3 5 si CO CD S1 •> CD 3 CD Q A CD'1 —^ CO CO CD 5-' 0 1 CD 3 I C3 CD CO Q . CD 3 CD CL CD O —\ o T3 zr 0) ? w co -a CD O CD' GO S? CD 0) cn o TI O cn .3 CD cn s? CD 01 cn O i cn 0) 3 CD cn s? CD 0 1 cn o i CO cu 3 CD J i cS z t7) C D 0) cn o •0 o cn 0) 3 CD cn Diversity analysis 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 ' i 0 ) , 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 (<x), Pielou (J'), Simpson index (1- ), and Taxonomic Diversity (8) indices. Of the remaining 3 indices, Species 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 1 s t / 2 n d year postharvest data, followed by a gradual decrease in 3 r d and 4/5 t h years. This trend occurred without significant difference for baited and control data in either index. The second trend observed in richness indices H ' i 0 , 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 2 n d year postharvest, falling to 101 Harvest Year Post 1 Post 2 Post 3 Post 4/5 Harvest Year g 0.3-% 0.2. Preharwst Post 1 Post 2 Post 3 Post 4/5 Harvest Year §—-§E S Preharvest Post 1 Post 2 Post 3 Harvest Year Preharvest Post 1 Post 2 Post 3 Post 415 Harvest Year Preharvest Post 1 Post 2 Post 3 Posl 4/5 Harvest Year tes Diversity (Brillouin; Spec Preharvest Post 1 Post 2 Post 3 Post 4/5 Harvest Year 8 i 0.8 Post 2 Post 3 Harvest Year Figure 3.3. Measu res of f lying beetle diversi ty f rom pheromone-ba i ted ( • ) and unbai ted ( • ) funnel t raps in Mature/Old growth nor thern inter ior Douglas-f i r ( F d 8 , >100 yrs). Preharvest th rough 4/5 season postharvest condi t ions. Pheromone t raps were bai ted wi th a f rontal in, M C O L , seudeno l lure speci f ic to Douglas-f i r beetle (oc = 0.05). 1.7a) Spec ies # (S), b) Margalef 's d, c) Pei lou's J ' , d) Bril louin, e) Shannon 's H', f) 1-Simpsons g) Fisher 's a, h) T a x o n o m i c divers i ty 8, i) Taxonomic d is t inctness 8*. 102 intermediate levels in 3 r a (98.80) and 4/5 t n years (88.20). The trend observed in control data was similar to that of pheromone-baited data. Single replicate data points for 1 s t and 2 n d 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 2 n d year postharvest, followed by a decrease to intermediate levels in 3 r d and 4/5 t h 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' i 0 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' 1 0 , & 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' 1 0 , 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 f lying Coleoptera statistically more or less abundant be tween pheromone-ba i ted and unbaited control sites in preharvest, 3 r d , and 4 / 5 t h season postharvest condi t ions. Forest habitat w a s mature to overmature interior Douglas-f i r undar at tack by the Douglas fir beetle prior to harvest ing. Pheromone-ba i ted t raps were baited with a frontal in, MCOL, seudenol lure for Douglas-f i r beet les (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 1 s t , 2 n d , 3 r d , and 4/5 t h 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. T rends in f ly ing Coleoptera t rapped in Douglas-f i r beetle p h e r o m o n e -bajted L indgren funnel t raps under preharvest and postharvest condi t ions. T rends are presented as the s u m of mean abundances /s i te for species wi th in a t rend. 354 spp fol lowed 1 of 4 t rends: Increasing m e a n abundance/s i te i ( 4 9 species) ; decreas ing m (42 spec ies) ; increasing then decreas ing ^ (253 spec ies) ; or decreas ing then increas ing >K (7 spec ies) f r o m preharvest through postharvest condi t ions. 103 spp (not shown) fo l lowed no d iscernable t rend. Spec ies with single occur rences (132) were omi t ted f r o m analysis. 10000 0.1 -I , 1 . 1 1 Preharvest Post 1 Post 2 Post 3 Post 4/5 Harvest Year Figure 3.5. Observed t rends in f lying Co leoptera t rapped in preharvest and postharvest condi t ions t rapped by unbai ted L indgren funnel t raps. T rends are presented as the s u m of m e a n abundances /s i te for species wi thin a t rend. 186 spp fo l lowed 1 of 4 t rends: Increasing m e a n abundance/s i te A(54 species) ; decreas ing a (17 spec ies) ; increasing then decreas ing £ (96 spec ies) ; or decreas ing then increasing J K (20 spec ies) f rom preharvest through postharvest condi t ions. A n addi t ional 97 spp (not shown) fo l lowed no d iscernable t rend. Spec ies with single occur rences (101) were omi t ted f rom 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 4/5 t h 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. Succes ive phases on the invertebrate communi ty exploit ing dead w o o d . Phase A - a shor t - term stage mostly with species feeding on bark, such as Scolyt idae, Cerambyc idae, and species living in their cavit ies, as wel l as parasi tes and predators. Phase B - a rather short s tage with species mainly living under bark and in the sur face layer of t imber, and species associated with fungi. Bark b e c o m e s loose and falls off the s tem. Phase C - a long term stage wh ich can take several decades , most ly with w o o d inhabit ing species. Phase D - A very long stage dur ing wh ich many w o o d inhabit ing species are replaced by species living under the shelter of decay ing logs. Note that much variat ion occurs depend ing on the tree spec ies and geograph ica l areas in quest ion (according to Ehnst rom, as presented in Hel iovaara and Va isanen 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 conifer 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 sphegeus b-a clerid predator on Dendroctonus spp. Cucujus claviceps c - 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 pines (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 ( b c ) . 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 1 s t season through 4/5 t h 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 4/5 t h 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 1 s t and 2 n d 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 Col lect ion 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, 1 s t , 2 n d , 3 r d , & 4/5 t h 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, 1 s t season postharvest conditions and later 3 r d and 4/5 t h 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 4/5 t h 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 f rom 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 3 r d and 4/5 t h 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 4/5 t h 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 i 1) frontalin (Fn) (1,5,-dimethyl-6,8-dioxabicyclo[3.2.1]octane), racemic i 1) 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. 165 Bibliography Allison, J. D., J. H. Borden, R. L. Mcintosh, P, Groot and R. Gries. 2001. Kairomonal response by four Monochamus species (Coleopter: Cerambycidae) to Bark Beetle Pheromones. J. Chem. Ecol. 27(4): 633-646. Anonymous. 1995. Forest Practices Code of British Columbia: Bark Beetle Management Guidebook. Ministry of Forests, Victoria, B.C. Pp. 58. Anonymous. 1996. Ecology and Management of Douglas-fir at the Northern Limits of its Range. Workshop Proceedings. October 7,8,9,1996. Fort St James, British Columbia. Anonymous. 1998. Species Inventory Fundamentals: Standards for Components of British Columbia's Biodiversity No .1. Resources Inventory Committee Victoria, B.C. (Digital copy: www.publications.qov.bc.ca). Pp. 119. Ascoli-Christensen, A., S. M. Salom and T. L. Payne. 1993. Olfactory receptor cell responses of Ips grandicollis (Eichhoff) (Coleoptera: Scolytidae) to intra- and interspecific behavioral chemicals. J. Chem. Ecol. 19(4): 699-712. Asquith, A. Lattin, J. D. and A. R. Moldenke. 1990. Arthropods: The invisible diversity. Northwest Environ. J. 6: 404-405. Atkins, M. D. 1975. On factors affecting the size, fat content and behavior of a Scolytid. Z. Angew. Entomol. 78: 209-218. Aukema, B. H., D. L. Dahlsten, and K. F. Raffa. 2000. Improved population monitoring of bark beetles and predators by incorporating disparate behavioral responses o semiochemicals. Environ. Entomol. 29(3): 618-629. Bakke, A. and T. Kvamme. 1981. Kairomone response in Thanasimus predators to pheromone components of Ips typographus. J. Chem. Ecol. 7(2): 305-312. Barnes, B.V., D. R. Zak, S. R. Denton and S. H. Spurr. 1998. Forest Ecology 4th Edition. John Wiley and Sons, NY. 744 pp. Berryman, A. A. 1986. Forest Insects: Principles and Practice of Population Management. Plenum Press, New York. Pp. 17-49. Billings, R. F. 1985. Southern pine bark beetles and associated insects: Effects of rapidly-released host volatiles on response to aggregation pheromones. Z. Angew. Entomol. 99: 483-491. Billings, R. F. and R. S. Cameron. 1984. Kairomonal responses of Coleoptera, Monochamus titillator (Cerambycidae), Thanasimus dubius (Cleridae), Temnochila virescens (Trogostidae), to behavioral chemicals of southern pine bark beetles (Coleoptera: Scolytidae). Environ. Entomol. 13(6): 1542-1548. 166 Birch, M. C. 1974. Introduction. In M. C. Birch (ed.). Pheromones. American Elsevier Publishing Company, NY. Pp 1-7. Birch, M. C. 1984. Aggregation in bark beetles, in W. J. Bell and R. T Carde (eds.). Chemical Ecology of Insects. Chapman and Hall, London Pp 331-353. Birch, M. C , P. Svihra, T. D. Payne, and J. C. Milller. 1980. Sequence of chemically mediated behavior on host tree colonization by four cohabitating species of bark beetles. J. Chem. Ecol. 6(2): 395-414. Borden, J. H. 1974. Aggregation pheromones in the Scolytidae. In M. C. Birch, ed., Pheromones. American Elsevier Publishing Company, NY. Pp 135-160. Borden J. H. 1994. Future of semiochemicals for the managment of bark beetle populations. USDA Forest Service, Pacific South West Research Station General Technical Report* 150 (FS-PSW-GTR-150). Pp. 5-10. Borden J. H. and L. J. Chong and B. S. Lindgren. 1990. Redundancy in the semiochemical message required to induce attack on lodgepole pine by the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae). Can. Entomol. 122 (10): 769-777. Borden, J. H., J. R. Handley, J. A. McLean, R.M. Silverstien, L. Chong, K. N. Slessor, B. D. Johnson and H. R. Schuler. 1980. Enantiomer-based specificity in pheromone communication by two sympatric Gnathotrichus species (Coleoptera: Scolytidae). J. Chem. Ecol. 6(2): 445-456. Borden J. H., G. Gries, G. Chong, L. J. Werner, R. A. Holsten, E. H. Weiser, E. A. Dixon, H. F. Cerezke. 1996. Regionally specific bioactivity of two new pheromones for Dendroctonus rufipennis (Kirby) (Coleoptera: Scolytidae). J. Appl. Entomol. 120(6): 321-326. Bormann, F. H. and G. E. Likens 1979. Pattern and Process in a Forested Ecosytem. Springer-Verlag, New York. Pp. 253. Bowers, W. W., and J. H. Borden. 1992. Attraction of Lasconotus intricatus Kraus. (Coloeptera: Colydiidae) to the aggregation pheromone of the four eyed spruce beetle Polygraphus rufipennis (Kirby) (Coleoptera: Scolytidae). Can Entomol. 124(1): 1-5. Bowers, W. W., J. H. Borden, and A. G. Raske. 1996. Bionomics of the four-eyed spruce beetle, Polygraphus rufipennis (Kirby) (Coleoptera: Scoyltidae) in Newfoundland II. Host colonization sequence. J. Appl. Entomol. 120(8): 449-461. Bright, D. E. 1976. The Insects and Arachnids of Canada, Part 2: The Bark Beetles of Canada and Alaska (Coleoptera: Scolytidae). Canada Department of Agriculture, Publication # 1576. Pp. 242. Burt, P. J. A., and D. E. Pedgely. 1997. Nocturnal insect migration: Effects of local winds. In Advances in Ecological Research, 27: 61-92. 167 Bury, R. B. and P. S. Corn. 1988. Douglas-fir Forests in the Oregon and Washington Cascades: Relation of the Herpetofauna to Stand Age and Moisture. In symposium proceedings, Management of Amphibians, Reptiles, and Small Mammals in North America. USDA RM General Technical Report #166: 11-22. Bush, G. L. and M. A. Hoy. 1984. Evolutionary processes in insects. In CB. Huffaker, and R. L. Rabb (eds.), Ecological Entomology. John Wiley and Sons, NY. Pp. 247-278. Byers J. A. 1992. Attraction of bark beetles, Tomicus piniperda, Hylurgops palliatus and Trypodendron domesticum and other insects to short-chain alcohols and monoterpenes. J. Chem. Ecol. 18(12): 2385-2402. Byers, J. A. 1993. Simulation and equation models of insect population control by pheromone-baited traps. J. Chem. Ecol. 19(9): 1939-1956. Byers, J. A. 1987. Interactions of pheromone component odor plumes of western pine beetle. J. Chem. Ecol. 13: 2143-2157., Byers J. A. 1995. Host tree chemistry affecting colonization in bark beetles. In R. T. Carte and W. J. Bell, (eds.). Chemical Ecology of Insects 2. Chapman and Hall, NY. Pp. 154-213. Byers, J. A., O. Anderbrant, and J. Lofqvist. 1989. Effective attraction radius: A method for comparing species attractants and determining densities of flying insects. J. Chem. Ecol. 15(2): 749-765. Carde, R. T., and T. C. Baker. 1984. Sexual communication with pheromones, in W. J. Bell and R. T Carde (eds.). Chemical Ecology of Insects. Chapman and Hall, London. Pp 355-383. Carpenter, S. E., M. E harmon, E. R. Ingham, R. G. Kelsey, J. D. lattin, and T. D. Schowalter. 1988. J. R. Soc. Edinburgh. 94B: 33-43. Caza, C. L. 1993. Woody Debris in the Forests of British Columbia: A review of the literature and Current Research. British Columbia Land Management Report number 78. Pp. 99. Chapman, J. A. 1955. Towards and insect ecology. Can. Entomol. 87: 172-177. Chapman, J. A. 1963. Field selection of different log odors by Scolytid beetles. Can. Entomol. 95(7): 673-676. Chapman, J. A. and J. M Kinghorn. 1955. Window flight traps for insects. Can Entomol. 87:46-47. 168 Chenier, J.V. R. and B. J. R. Pilogene 1989. Field responses to certain forest Coleoptera to conifer monoterpenes and ethanol. J. Chem. Ecol. 15(6): 1729-1745. Church, R., R. Gerrard, A Hollander, and D. Stoms. 2000. Understanding the tradeoffs between site quality and species presence in reserve site selection. For. Sci. 46(2): 157-167. Clarke K. R., and R. M. Warwick. 1998. A taxonomic distinctness index and it's statistical properties. J. Appl. Ecol. 35: 523-531. Clarke K. R., and R. M. Warwick. 1999. The taxonomic distinctness measure of biodiversity: Weighting of step lengths between hierarchical levels. Mar. Ecol. Prog. Ser. 184: 21-29. Collins, B. S., K. P. Dune and T. S. Pickett. 1985. Responses of forest herbs to canopy gaps. In S. T. A. Pickett and P. S. White, eds. The ecology of natural disturbance and patch dynamics. Academic Press, Inc. Pp 218-234. Dajoz, R. 2000. Insects and Forests: The Role and Diversity of Insects in the Forest Environment. Intercept Ltd. Paris, France. Pp. 668. De Leon, D. 1934. An Annotated List of the parasites, Predators, and Other Associated Fauna of the Mountain Pine Beetle In Western White Pine and Lodgepole Pine. Can. Entomol. 66(3):51-61. Delorme, J. D. and T. L. Payne. 1990. Antennal olfactory responses of black turpentine beetle, Dendroctonus terebrans (Olivier), to bark beetle pheromones and host volatiles. J. Chem. Ecol. 16(4): 1321-1329. Deyrup, M. A. and R. I. Gara. 1978. Insects Associated with Scolytidae (Coleoptera) in Western Washington. Pan. Pac. Entomol. 54: 270-282. Dickens, J. C, T. L. Payne, L. C. Ryker and J. A. Rudinsky. 1984. Single cell response, of the Douglas-fir beetle Dendroctonus pseudotsugae Hopkins (Coleoptera: Scolytidae), to pheromones and host odors. J. Chem. Ecol. 10(4): 583-600. Dickens, J. C, T. L. Payne, L. C. Ryker and J. A. Rudinsky. 1985. Multiple acceptors for pheromonal enatiomers on single olfactory cells in the Douglas-fir beetle Dendroctonus pseudotsugae Hopk. (Coleoptera: Scolytidae). J. Chem. Ecol. 11(10): 1359-1370. Dixon, W. N. and T. L Payne. 1980. Attraction of entomophagous and associate insects of the southern pine beetle- to beetle and host tree-produced volatiles. J. Georgia Entomol. Soc. 15(4): 378-389. Dubbel, V., K. Kerck, M. Short and S. Mangold. 1985. Influence of trap color on the efficiency of bark beetle pheromone traps. Z. Angew. Entomol. 99: 59-64. 169 Dyer, E. D. A. 1973. Spruce beetle aggregated by the synthetic pheromone frontalin. Can J. For. Res. 3: 486-494. Edmonds, R. L. and A. Eglitis. 1989. The role of the Douglas-fir beetle and wood borers in the decomposition of and nutrient release from Douglas-fir logs. Can. J. For. Res. 19: 853-859. Elkinton, J. S., and R. T. Carde. 1984. Odor Disperson, in W. J. Bell and R. T Carde (eds.). Chemical Ecology of Insects. Chapman and Hall, London. Pp 73-91. Farkas, S. R. and H. H. Shorey. 1974. Orientation to a distant pheromone source. In M. C. Birch, ed., Pheromones. American Elsevier Publishing Company, NY. Pp 81-95. Fletchmann, c. A. H., A. L. T. Ottati and c. W. Berisford. 2000. Comparison of four trap types for ambrosia beetles (Coleoptera: Scolytidae) in Brazilian Eucalyptus stands. J. Econ. Entomol. 93(6): 1701-1707. Furniss, M. M., G. E. Daterman, L. N. Kline, M. D. McGregor, G. C. Trostle, L F. Pettinger and J. A. Rudinsky. 1974. Effectiveness of the Douglas-fir beetle antiaggregative pheromone methylcyclohexenone at three concentrations and spacings around felled host trees. Can. Entomol. 106: 381-392. Furniss, M. M., B. H. Baker and B. B Hostetler. 1976. Aggregation of spruce beetles (Coleoptera) to seudenol and repression of attraction by methylcyclohexenone in Alaska. Can. Entomol. 108(12): 1297-1302. Furniss, R. L. and V. M. Carolin. 1977. Western Forest Insects. USDA Miscellaneous Publication # 1339. Pp. 654. Gara, R. I., R. A. Werner, M.C. Whitmore, and E. H. Hosten. 1995. Arthropod associates of the spruce beetle, Dendroctonus rufipennis (Kirby) (Col., Scolytidae) in spruce stands of south-central and interior Alaska. J. Appl. Entomol. 119: 585-590. Gast, S. J., M. W. Stock, and M. M. Furness. 1993. Physiological odors affecting attraction of Ips pini (Coleoptera, Scolytidae) to host odor or natural male pheromone in Idaho. Ann. Entomol. Soc. Am. 86(4): 417-422. Greenwood, M. E. and J. H. Borden. 1999. Co-baiting for spruce beetles, Dendroctonus rufipennis, and western bark beetles, Dryocetes confuses (Coleoptera: Scolytidae). Can. J. For. Res. 30: 50-58. Gries G. 1992. Ratios of geometrical and optical isomers of pheromones: Irrelevant or Important in Scolytids. J. Appl. Entomol. 114(2): 240-243. Grosman, D. M., S. M. Salom, F. W. Ravlin, and R. W. Young. 1997. Geographic and gender differences in semiochemicals in emerging adult southern pine beetle (Coleoptera: Scolytidae). Ann. Entomol. Soc. Am. 90(4): 438-446. 170 Guthrie, S. and H. Wieser. 1994. Douglas-fir beetle pheromone baiting 1994 in the Fort St James Forest District. Report to the B.C. Ministry of Forests, Fort St James Forest District, Fort St James, B.C. 41 pp. Guthrie, S. and H. Wieser. 1997. Douglas-fir beetle pheromone baiting 1997 in the Fort St James Forest District. Report to the B.C. Ministry of Forests, Fort St James Forest District, Fort St James, B.C. 31 pp. Haack, R. A. and J. W. Byler 1993. Insects and pathogens: Regulators of forest ecosytems. J. For. 91(9): 32-37 Halme E. and J. Niemela. 1993. Carabid beetles in fragments of coniferous forests. Ann. Zool. Fenn. 30: 17-30. Hammond, H. E. J. 1997. Arthropod biodiversity from Populus coarse woody material in nort-central Alberta: A review of Taxa and collection methods. Can. Entomol. 126:1009-1033. Harmon, M. E., J. F. Franklin, F. J. Swanson, P. Sollins, S. V. Gregory, J. D. Lattin, N. H. Anderson, S. P. Cline, N. G. Alumen, J. R. Sedwell, G. W. leinkaemper, K. Cromack Jr., and K. W. Cummins. 1986. Ecology of coarse woody debris in temperate ecosystems. Rec. Adv. Ecol. Res. 15: 135-305. Harris, M. O. and S. P. Foster. 1995. Behavior and integration. In R. T. Carte and W. J. Bell, eds., Chemical Ecology of Insects 2. Chapman and Hall, NY. Pp. 3-46. Hedden R., J. P Vite, and K. Mori. 1976. Synergistic effect of a pheromone and a kairomone on host selection and colonisation by Ips avulsus. Nature 261 (June 24): 696-697. Hedden, R. L. and G. B. Pitman. 1978. Attack density regulation: A new approach to the use of pheromones in Douglas-fir beetle population management. J. Econ. Entomol. 71: 633-637. Heikkenen, H. J., and B. F. Hrutfiord. 1965. Dendroctonus pseudotsugae: A hypothesis regarding its primary attractant. Science 150: 1457-1459. Heliovaara, K. and R. Vaisanen. 1984. Effects of modern forestry on northwestern European forest invertebrates: A synthesis. Acta For. Fenn. 198: 5-29. Hermann, R. K. and D. P. Lavender. 2001. Pseudotsuga menziesii (Mirb) Franco: Douglas-fir www.na.fs.fed.us/spfo/pubs/silvics_manualA/olume_1/pseudotsuga/menziesii. htm Herms, D. A., R. A. Haack, and B. D. Ayers. 1991. Variation in semiochemical-mediated prey-predator interaction: Ips pini (Scolytidae) and Thanasimus dubius (Cleridae). J. Chem. Ecol. 17(8): 1705-1714. 171 Huber D. P. W., R. Gries, J. H. Borden and h. D. Pierce. 2000. A survey of antennal responses by five species of coniferophagous bark beetles (Coleoptera: Scolytidae) to bark volatiles of sec specis of angiosperm trees. Chemoecology 10: 103-113. Huffaker, C. B., D. L. Dahlsten, D. H. Jansen, and G. G. Kennedy. 1984. Insect influences in the regulation of plant communities. In CB. Huffaker, and R. L. Rabb, eds., Ecol. Entomol. John Wiley and Sons, NY. Pp. 659-691. Hutcheson, J. 1990. Characterization of terrestrial insect communities using quantified, Malaise-trapped Coleoptera. Ecol. Entomol. 15: 143-151. Jakus, R. 1998. Patch level variation on bark beetle attack (Col., Scolytidae) on snapped and uprooted trees in Norway spruce primeval natural forest in endemic condition: Effects of host and insulation. J. Appl. Entomol. 122: 409-421. Jonsson, B. G., and M. Jonsell. 1999. Exploring potential biodiversity indicators in boreal forests. Biodiv. and Conserv. 8: 1417-1433. Kangas, J. and J. Kuusipalo. 1993. Integrating biodiversity in forest management planning and decision-making. For. Ecol. and Manag. 61: 1-15. Kareiva, P. 1986. Patchiness, dispersal and species interactions: consequences for communities of herbivorous insects. In J. Diamond and T. J Case, eds. Community Ecology. Harper and Row, NY. Pp 192-206. Karr, J. R. and K. E. Freemark. 1985. Disturbance and vertebrates: an integrated approach. In S. T. A. Pickett and P. S. White, eds. The ecology of natural disturbance and patch dynamics. Academic Press, Inc. Pp 153-168. Kinzer, G. W., A. F. Fentiman, Jr., R. L. Flotz and J. A. Rudinsky. 1971. Bark beetle attractants: 3-methyl-2-cyclohexen-1-one isolated from Dendroctonus pseudotsugae. J. Econ. Entomol. 970-971. Kirk, W. D. 1984. Ecologically selective coloured traps. Ecol. Entomol. 9: 35-41. Kitching, R. L, D. Li and N. E. Stork. 2001. Assessing biodiversity 'sampling packages': How similar are arthropod assemblages in different tropical rainforests? Biodiv. and Conserv. 10: 793-813. Knight, F. B. and H. J. Heikkenen. 1980. Principles of Forest Entomology, Fifth Edition. McGraw-Hill Inc. New York. Pp. 331-409. Kogan, M. and J. D. Lattin. 1993. Insect conservation and pest management. Biodiv. and Conserv. 2: 242-257. Kohnle, U. and J. P. Vite. 1984. Bark beetle predators: Strategies in the olfactory perception of prey species by clerid and trogostid beetles. Z. Angew. Entomol. 98: 504-508. 172 Krebs, C. (1994). Ecology: The Experimental analysis of Distribution and Abundance. Harper Collins College Publishers. 801 pp. Lanier, G. N. 1970. Sex pheromones: Abolition of specificity in hybrid bark beetles. Science, 169: 71-72. Lanier, G. N. and D. L. Wood. 1975. Specificity of response to pheromones in the genus Ips (Coleoptera: Scolytidae). J. Chem. Ecol. 1(1): 9-23. Lattin, J. D. 1993. Arthropod diversity and conservation in old-growth northwest forests. Am. Zool. 33: 578-587. Lautenschlager, R. A. 1997. Biodiversity is dead. Wild. Soc. Bull. 25(3): 679-685. Lavallee, S. L. 1999. Changes in the carabid community of the Sicamous Creek research site in response to prescribed logging practices. M. Sc. Thesis. University of British Columbia, Vancouver, B. C. 72 pp. Lehmkuhl, D. M., H. V. Danks, V. M. Behran-Pelletier, D. J. Larson, D. M. Rosenberg, and I. M. Smith. 1984. Recommendations for the appraisal of environmental disturbance: Some general guidelines, and the value and feasibility of insect studies. Supplement to The Entomological Society of Canada Bulletin 16(3): 8 pp. Lejeune, R. R., L. H. McMullen and M. D. Atkins. 1961. The influence of logging on Douglas-fir beetle populations. For. Chron. 37: 308-314. Lenski, R. E. 1982. The impact of forest cutting on the diversity of ground beetles (Cloeoptera: Carabidae) in the southern Appalachians. Ecol. Entomol. 7:385-390. Lessard, E. D., and J. M. Schmid. 1990. Emergence, attack densities, and host relationships for the Douglas-fir beetle (Dendroctonus pseudotsugae Hopkins) in northern Colorado. Great Basin Nat. 50(4): 333-338. Lewis, K. J., and B. S. Lindgren. 2000. A conceptual model of biotic disturbance ecology in the central interior of B.C.: How forest management can turn Dr. Jekyll into Mr. Hyde. For. Chron. 76(3): 433-443. Libbey, L. M., A. C. Oehlschlager and L. C. Ryker. 1983. 1-Methylcyclohex-2-en-ol as an aggregation pheromone of Dendroctonus pseudotsugae. J. Chem. Ecol. 9(2): 1533-1541. Libbey, L. M., L. C. Ryker, and K. L. Yandell. 1985. Laboratory and field studies of volatiles released by Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae). Z. Angew. Entomol. 99:381-392. Lindgren, B. S. 1983. A multiple funnel trap for Scolytid beetles (Coleoptera). Can. Entomol. 115: 299-302. 173 Lindgren, B. S. 1992. Attraction of Douglas-fir beetle, spruce beetle and a bark beetle predator (Coleoptera: Scolytidae and Cleridae) to enantiomers of frontalin. J. Entomol. Soc. Brit. Columbia 89: 13-17. Lindgren, B. S., G. Gries, H. D. Pierce, Jr., and K. Mori. 1992. Dendroctonus pseudotsugae Hopkins (Coleoptera: Scolytidae): Production of and response to enantiomers of 1-methycyclohex-2-en-1-ol. J. Chem. Ecol. 18(7): 1201-1207. Lindroth, C. H. 1968. The ground-beetles (Carabidae excluding Cicindelidae) of Canada and Alaska. Part 5. Opusc. Entomol., Supplementum 33: 649-944. Liu, Y. and J. A. McLean. 1993. Observations on the biology of the ambrosia beetle Gnathotrichus retusus (Leconte) (Coleoptera: Scolytidae). Can. Entomol. 125: 73-83. Lundquist, J. E. 1995. Pest interactions and canopy gaps in ponderosa pine stands in the Black Hills, South Dakota, USA. For. Ecol. and Manag. 74: 37-48. MacDonald, R. C. and T. W. Kimmerer. 1991. Ethanol in the stems of trees. Physiol. Plant. 82(4): 582-588. Mackinnon, A., J. Polar, and R. Coupe. 1992. Plants of Northern British Columbia. Lone Pine Publishing, Edmonton, Alberta. Pg. 20. Maddock, A., and M. A. Du Plessis. 1999. Can species data only be appropriately used to conserve biodiversity? Biodiv. and Conserv. 8: 603-615. Magurran, A. E. 1988. Ecological Diversity and its Measurement. Princeton University Press. 167 pp. Marshall, S. A; R. S. Anderson, R. E. Roughly, V. Behan-Pelletier, and V. Danks. 1994. Terrestrial Arthropod Diversity: Planning a Study and Recommended Sampling Techniques. A Brief Prepared for the Biological Survey of Canada (Terrestrial Arthropods). Bull. Entomol. Soc. Canada. 26(1), supplement, 33pp. Maser, C, S. P. Cline, K. Cromack Jr., J. M. Trappe and E. Hansen. 1988. what we know about large trees that fall to the forest floor. In Maser, C, R. F. Tarrant, J. M. Trappe, and J. F. Franklin Editors. From the forest to the sea: A story of fallen trees. USDA, PNW General Technical Report # 229: 25-45. May, R. E. 1976. Patterns in multi-species communities. As found in Theoretical Ecology. Principles and Applications. R. May - Ed. W.B. Sanders Company, pp. 142-162. McDonald R. C, and T. W. Kimmerer 1991. Ethanol in the stems of tress. Physiologia Panatarum 82: 582-588. McDowell, J. K. 1998. Response of Carabid Species and Assemblages to Forest Practices of British Columbia in Englelmann Spruce - Subalpine Fir and 174 Interior Cedar-Hemlock Forests. M. Sc. Thesis, University of British Columbia, Vancouver, B.C. 106 pp. McLean, J. A., A. Bakke, and H. Niemeyer. 1987. An evaluation of three traps and two lures for the ambrosia beetle Trypodendron Linneatum (Oliv.) (Coleoptera: Scolytidae) in Canada, Norway and West Germany. Can. Entomol. 119:273-280. McLeod, J. M. 1980. Forests, disturbances and insects. Can. Entomol. 112: 1185-1192. McMullen, L. H. and M. D. Atkins. 1962. On the flight and host selection of the Douglas-fir beetle, Dendroctonus pseudotsugae Hopk. (Coleoptera: Scolytidae). Can. Entomol. 94: 1309-1324. Meidinger, D. and J. Pojoar. 1991. Ecosystems of British Columbia. Research Branch, Ministry of Forests. Victoria, B.C. 330 pp. Miller, J. C. 1993. Insect natural history, multi-species interactions and biodiversity in ecosystems. Biodiv. and Conserv. 2: 233-241. Milton, J. S. 1992. Statistical Methods in the Health and Biological Sciences. McGraw Hill Inc. NY. Pp. 433-437. Montgomery, M. E. and P. M. Wargo. 1983. Ethanol and other host-derived volatiles as attractants ot beetles that bore into hardwoods. J. Chem. Ecol. 9(2): 181-190. Muirhead-Thomson, R. C. 1991. Trap responses of flying insects. Academic Press, NY. 287 pp. Mustaparta, H. 1984. Olfaction, in W. J. Bell and R. T Carde (eds.). Chemical Ecology of Insects. Chapman and Hall, London. Pp 37-72. Negron J. F. 1998. Probability of infestation and extent of mortality associated with the Douglas-fir beetle in the Colorado front range. For. Ecol. & Manag. 107(1-3): 71-85. Negron, J. F., W. C. Schaupp Jr., K. E. Gibson, J. Anhold, D. Hansen, R. Thier, and P. Mocettini. 1999. Estimating extent of mortality associated with Douglas-fir beetle in the central and northern Rockies. West. J. Appl. For. 14(3): 121-127. Niemela, J., D. Langor, and J. R. Spence. 1993. Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae). Conserv. Biol. 7(3): 551-559. Niemela, J., J. R. Spence, D. Langor, H. Tukia and Y. Haila. 1992. Logging and boreal ground-beetle assemblages on two continents: Implications for conservation. In Gaston, K. J., et al. (eds.), Perspectives in Insect Conservation, Intercept Publishers Ltd., Andover, U. K. 175 Nilsson, S. G. and R. Baranowski. 1997. Habitat predictability and the occurrence of wood beetles in old growth beech forests. Ecography 20: 491-498. Nilsson, S. G., U. Arup, R. Baranowski, and S. Ekman. 1994. Tree-dependant lichens and beetles as indicators in conservation forests. Conserv. Biol. 9(5): 12-8-1215. Noss, R. F. 1990. Indicators for monitoring biodiversity: A hierarchical approach. Conserv. Biol. 4(4): 355-364. Oliver, C. D. and B. C. Larson. 1990. Forest Stand Dynamics. McGraw Hill Inc. Pp. 117-126. Paine, T. D., J. G. Millar, C. C. Hanlon, and J. S. Hwang. 1999. Indentification of semiochemicals associated with Jeffrey Pine Beetle, Dendroctonus jeffreyi. J. Chem. Ecol 25(3): 433-453. Paulson, L. C. 1995. Monitoring and dynamics of a Douglas-fir beetle outbreak in Jasper National Park, Alberta. J. Entomol. Soc. Brit. Columbia 92: 17-23. Payne, T. L. 1974. Pheromone perception. In M. C. Birch (ed). Pheromones. American Elsevier Publishing Company, NY. Pp 35-61. Payne, T. L, J. E. Coster, J. V. Richerson, L. J. Edson, and E. R. Hart. 1978. Field response of the southern pine beetle to behavioral chemicals. Environ. Entomol. 7: 578-582. Payne, T. L., J. C. Dickens and J. V. Richerson. 1984. Insect predator-prey coevolution via enantiomeric specificity in a Kairomone-pheromone system. J. Chem. Ecol. 10(3): 487-492. Payne, T. L., R. F. Billings, J. D. Delorme, N. A. Andryszak, J. Bartels, W. Francke, and J. P. Vite. 1987. Kairomonal-pheromonal system in the back turpentine beetle, Dendroctonus terebrans. J. Appl. Entomol. 103(1): 15-22. Payne, T. L., N. A. Andryszak, H. Wieser, E. A. Dixon, N Ibrahim, and J. Coers. 1988. Antennal olfactory and behavioral response of the southern pine beetle Dendroctonus frontalis, to analogs of its aggregation pheromone frontalin. J. Chem. Ecol. 14(4): 1217-1225. Peck, R. W., A. Equihua-Martinez, and D. W. Ross. 1997. Seasonal flight patterns of bark and ambrosia beetles (Coleoptera: Scolytidae) in northeastern Oregon. Pan-Pacific Entomol. 73(4): 204-212. Perry, D. A. 1994. Forest Ecosystems. The Johns Hopkins University Press Ltd. Baltimore. 649pp. Phillips, T. W., A. J. Wilkening, T. H. Atkinson, J. L. Nation, R. C. Wilkenson and J. L. Flotz. 1988. Synergism of turpentine and ethanol as attractants for certain pine-infesting beetles (Coleoptera). Environ. Entomol. 17(3): 456-462. 176 Pielou, E. C. 1969. An Introduction to Mathematical Ecology. Wiley-lnterscience, New York. Pp. 222-236. Pitman, G. B. and J. P. Vite. 1970. Field response of Dendroctonus pseudotsugae (Coleoptera: Scolytidae) to synthetic Frontalin. Ann. Entomol. Soc. Am. 63(2): 661-664. Pitman, G. B. and J. P. Vite. 1974. Biosynthesis of Methylcyclohexenone by male Douglas-fir beetle. Environ. Entomol. 3(5): 886-887. Pitman, G. B., R. L. Hedden, and R. I. Gara. 1975. Synergistic effects of ethyl alcohol on the aggregation of Dendroctonus pseudotsugae (Coleoptera: Scolytidae) in response to pheromones. Z. Angew. Entomol. 78: 203-208. Plummer, E. L, T. E. Stewart, K. Byrne, G. T. Pearce and R. M. Silverstein. 1976. Determination of the enantiomeric composition of several insect pheromone alcohols. J. Chem . Ecol. 3(2): 307-331. Prenzel, B. G., W. G. Laidlaw, and H. Wieser. 1999. Within-tree dynamics of mass attack by the Dendroctonus pseudotsugae (Coleoptera; Scolytidae) on its host. Can. Entomol. 131(5): 635-643. Pureswaran, D. S., R. Gries, J. H. Borden and H. D. Pierce Jr. 2000. Dynamics of pheromone production and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae). Chemoecology. 10: 153-168. Raffa, K. F. 1991. Temporal and spatial disparities among bark beetles, predators, and associates responding to synthetic bark beetle pheromones: Ips pini (Coleoptera; Scolytidae) in Wisconsin. Environ. Entomol. 20(6): 1665-1679. Raffa K. F. 2001. Mixed messages across multiple trophic leves: The ecology of bark beetle chemical communication systems. Chemoecology 11: 49-65. Raffa, F., and K. D. Klepzig. 1989. Chiral escape of bark beetles from predators responding to a bark beetle pheromone. Oecologia 80: 566-569. Rankin, L. H. and J. Borden. 1991. Competitive interactions between mountain pine beetle and the Pine engraver in Lodge pole pine. Can. J. For. Res. 21: 1029-1036. Renwick, J. A. A., and P. R. Hughes. 1975. Oxidation of unsaturated cyclic hydrocarbons by Dendroctonus frontalis. Insect Biochem. 5(4): 459-463. Ringold, G. B., P. J. Gravelle, D. Miller, M. M. Furniss and M. D. McGregor. 1975. Characteristics of Douglas-fir Beetle Infestation in Northern Idaho Resulting From Treatment with Douglure. USDA Forest Service Research Note lnt-189. January, 1975. 177 Roling, M. P. and W. H. Kearby. 1975. Seasonal flight and vertical distribution of Scolytidae attracted to ethanol in an oak-Hickory forest in Missouri. Can. Entomol. 107: 1315-1320. Ross D. W. and G. E. Daterman. 1994. Reduction of Douglas-fir beetle infestation of high-risk stands by antiaggregation and aggregation pheromones. Can. J. For. Res. 24: 2184-2190. Ross D. W. and G. E. Daterman. 1995. Response of Dendroctonus pseudotsugae (Coleoptera: Scolytidae) and Thanasimus undatulus (Coleoptera: Cleridae) to traps with different semiochemicals. For. Entomol. 88(1): 106-111. Ross D. W. and G. E. Daterman. 1997. Using pheromone-baited traps to control the amount and distribution of tree mortality during outbreaks of the Douglas-fir beetle. For. Sci. 43(1): 65-70. Ross D. W. and G. E. Daterman. 1998. Pheromone-baited traps for Dendroctonus pseudotsugae (Coleoptera: Scolytidae): Influence of selected release rates and trap designs. J. Econ. Entomol. 91(2): 500-506. Rudinsky, 1966a. Scolytid beetles associated with Douglas-fir: Response to terpenes. Science 152: 218-219. Rudinsky, 1966b. Host selection and invasion by the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, in coastal Douglas-fir forests. Can. Entomol. 98: 98-111. Rudinsky, J. A. and O. Zethner-Moller. 1967. Olfactory responses of Hylastes nigrinus (Coleoptera: Scolytidae) to various host materials. Can. Entomol. 99: 911-916. Rudinsky, J. A., M. E. Morgan, L. M. Libbey, and T. B. Putnam. 1977. Limonene released by the Scolytid beetle Dendroctonus pseudotsugae. Z. Angew. Entomol. 82(4): 376-380. Salom, S. M. and J. A. McLean. 1991. Flight behavior of scolytid beetle in response to semiochemicals at different wind speeds. J. Chem. Ecol. 17(3): 647-661. Schlyter, F. 1992. Sampling range, attraction range, and effective attraction radius: Estimates of trap efficiency and communication distance in coleopteran pheromone and host attractant systems. J. Appl. Entomol. 114: 439-454. Schowalter, T. D. 1981. Insect herbivore relationship to the state of the host plant: biotic regulation of ecosystem nutrient cycling though ecological succession. Oikos 37(1): 126-130. Schowalter, T. D. 1985. Adaptations of insects to disturbance. In S. T. A. Pickett and P. S. White (eds). The ecology of natural disturbance and patch dynamics. Academic Press, Inc. Pp 235-252. 178 Schowalter, T. D., J. D. Lattin, R. G. Kelsey, S. E. Carpenter, E. R. Ingham, M. E. Harmon, and A. R. Moldenke. 1988. Insect mediated community development and decomposition in conifer logs in the Pacific Northwest. Proceedings of the XVIII International Congress of Entomology. Vancouver, British Columbia, Canada. July 3-9, 1988. Schlyter, F. 1992. Sampling range, attraction range, and effective attraction radius: Estimates of trap efficience and communication distance in conleopteran pheromone and lost attractant systems. J. Appl. Entomol. 114: 439-454. Scudder, G. G. E. 1996. Terrestrial and Freshwater Intertebrates of British Columbia: Priorities for Inventory and Descriptive Research. British Columbia , Ministry of Forests Research Program, Victoria, B. C. Pp. 205. Seip, D. 1996. The projected impacts of different biodiversity emphasis options on some forest bird species in the Sub-Boreal Spruce (SBS) zone. Forest Research Note # PG-04. Prince George Region. August 1996, 4pp. Setter, R. R., and J. H. Borden. 1992. Response by the striped ambrosia beetle, Trypodendron linneatum (Olivier), to the bark beetle pheromone, frontalin. Can. Entomol. 124: 559-560. Setter, R. R., and J. H. Borden. 1999. Bioactivity and efficacy of MCOL, and seudenol as potential attractive bait components for Dendroctonus rufipennis (Coleoptera: Scolytidae). Can. Entomol. 131: 251-257. Seybold, S. J. 1993. Role of chirality in olfactory-directed behavior: Aggregation of pine engraver seetles in the genus Ips (Coleoptera: Scolytidae). J. Chem. Ecol. 19(8): 1809-1831. Shore, T. L. and J. A. McLean. 1984. The effect of trap height of pheromone-baited traps on catches of the ambrosia beetle, Trypodendron linneatum. J. Entomol. Soc. Brit. Columbia. 81: 17-18. Shore, T. L., L. Safranyik, W. G. Riel, M. Ferguson, and J. Castonguay. 1999. Evaluation of factors affecting tree and stand susceptibility to the Douglas-fir beetle (Coleoptera: Scolytidae). Can. Entomol. 131(6): 831-839. Shore, T. L., P. M. Hall and T. F. Maher. 1990. Grid baiting of spruce stands with frontalin for pre-harvest containment of the spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Scolytidae). J. Appl. Entomol. 109(3): 315-319. Shore, T. L, W. G. Riel and L. Safranyik. 1996. A decision support system for the mountain pine beetle in lodgepole pine stands. In Shore, T. L. and D. A. Maclean (eds.), Decision Support Systems in Forest pest management. Proceedings of a workshop at the joint meeting of the entomological societies of Canada and British Columbia, October 17, 1995, Victoria, B. C, Canada. Canada-British Columbia Forest Resource Development Agency, Report # 260. Pp. 25-30. 179 Smetana, A. 1971. Revision of the tribe Quediini of America north of Mexico (Coleoptera: Staphylinidae) Mem. Entomol. Soc. of Canada 79: 107-109. Smith, D. B. and M. K. Sears. 1982. Mandibular structure and feeding habits of three morphologically similar coleopterous larvae: Cucujus claviceps (Cucujidae), Dendroides canadensis (Pyrochroidae), and Pytho depressus (Salpingidae). Can. Entomol. 114(1): 173-175. Smith, M. T., T. L. Payne and M. C. Birch. 1990. Olfactory based behavioral interactions among 5 species in the southern bark beetle group. J. Chem. Ecol. 16(12): 3317-3329. Southwood, T. R. E., V. K. Brown and P. M. Reader. 1979. The relationship of plant and insect diversities in succession. Biol. J. Linn. Soc. 12: 327-348. Stephen, F. M., and D. L. Dahlston. 1976. The arrival sequence of the athropod complex following attack by Dendroctonus brevicomis (Coleoptera: Scolytidae) in ponderosa pine. Can. Entomol. 108: 283-304. Su, J. C. and S. A. Woods. 2001. Importance of sampling along a vertical gradient to compare the insect fauna in managed forests. Environ. Entomol. 30(2): 400-408. Thier, R. W. and J. C. Weatherby. 1991. Mortality of Douglas-fir after two semiochemical baiting treatments for Douglas-fir beetle (Coleoptera: Scolytidae). J. Econ. Ent. 84(3): 962-964. Thier, R. W. and S. Paterson. 1997. Mortality of Douglas-fir after operational semiochemical baiting for Douglas-fir beetle (Coleoptera: Scolytidae). West. J. Appl. For. 12(1): 16-20 Tilden, P. E., W. D. Bedard, D. L. Wood, K. Q. Lindahl and P. A. Rauch. 1979. Trapping the western pine beetle at and near a source of synthetic attractive pheromone: Effects of trap size and position. J. Chem. Ecol. 5(4): 519-531. Toman M. A. and P M. Ashtom 1996. Sustainable forest management: A review article. For. Sci. 42(3); 366-377. Trueman, J. W. and P. S. Cranston. 1997. Prospects for the rapid assessment of terrestrial invertebrate biodiversity. Mem. Natl. Mus. Victoria 56(2): 349-354. Turchin, P. and F. J. Odendaal. 1996. Measuring the effective sampling area of a pheromone trap for monitoring population density of southern pine beetle (Coleoptera: Scolytidae). Environ. Entomol. 25(3): 582-588. Vaisanen, R., O. Bistrbm and K. Heliovaara. 1993. Sub-cortical Coleoptera in dead pines and spruces: is primeval species composition maintained in managed forests? Biodiv. and Conserv. 2: 95-113. 180 Van Kooten, G. C. 1994. Economics of Biodiversity and Preservation in Forestlands in British Columbia. Forest Resource Development Agency (FRDA) Report # 112. Pp. 35. Vite, J. P. and E. Baader. 1990. Present and future use of semiochemicals in pest management of bark beetles. J. Chem. Ecol. 16(11): 3031-3037. Warwick, R. M., and K. R. Clarke. 1995. New 'biodiversity' measures reveal a decrease in taxonomic distinctness with increasing stress. Mar. Ecol. Prog. Ser. 129: 301-305. Warwick, R. M., and K. R. Clarke. 1998. Taxonomic distinctness and environmental assessment. J. Appl. Ecol. 35: 532-543. Werner, R. A. 2002. Effect of Ecosystem disturbance on diversity of bark and wood-boring beetles (Coleoptera: Scolytidae, Buprestidae, Cerambycidae) in white spruce (Picea Glauca (moench) Voss) ecosystems of Alaska. USDA Forest Service, Pacific Northwest Research Station, Research Paper #546 (PNW-RP-546). Pp. 15. Werner, R. A. and E. H. Holsten. 1984. Scolytidae associated with felled white spruce in Alaska. Can. Entomol. 116: 465-471. Werner, R. A., M. M. Furniss, L. C. Yarger, and T. Ward. 1981. Effects on eastern larch beetle of its natural attractant and synthetic pheromones in Alaska. USDA Forest Service, Pacific Northwest Research Station, Research Note #371 (PNW-RN-371). Pp. 7 Werner, S. M. and K. F. Raffa. 2000. Effects of forest management practices on the diversity of ground-occurring beetles in mixed northern hardwood forests of the Great Lakes Region. For. Ecol. Manag. 139: 135-155. White, R. A., M. Agosin, R. T. Franklin and J. W. Webb. 1980. Bark beetle pheromones: Evidence for physiological synthesis mechanisms and their ecological implications. Z. Angew. Entomol. 90: 255-274. Wiens, J. A., J. F. Addicott, T. J. Case and J. Diamond. 1986. Overview: The importance of spatial and temporal scale in ecological investigations. In J. Diamond and T. J Case (eds). Community Ecology. Harper and Row, New York. Pp. 145-153. Wieser, H. and E. A. Dixon. 1992. Douglas-fir beetle pheromone application 1992 in the Invermere Forest District. Report to the B.C. Ministry of Forests, Invermere Forest District, Invermere, B.C., and Crestbrook Forest Industries Ltd., Cranbrook, B.C. Wigglesworth, V. B. 1984. Insect Physiology. Chapman and Hall, NY. Pp. 154-184. Wilkenson A. T. 1963. Wireworms of cultivated land. Proc. Entomol. Soc. Brit. Columbia. 60: 3-17. 181 Winchester, N. N. and G. E. E. Scudder. 1993. Methodology for Sampling Terrestrial Arthropods in British Columbia. Min. Environ., Lands and Parks, Victoria, B. C. RIC publication. Pp 32. Wood, D. L. 1982. The role of pheromones, kairomones and allomones in the host selection and colonization behavior of bark beetles. Ann. Rev. Entomol. 27:411-416. Wright, R. H. 1958. The olfactory guidance of flying insects. Can Entomol. Feb: 81-88. Zahradnik, P. 1995. Evaluation of non-target catches of beetles using the pheromone preparation of Chalcoprax in protective measures against the Pine bark beetle - Pityogenes chalcographus (Coleoptera, Scolytidae). Zapravy-Lesnickeho-Vyzkumu 40:2, 13-19. 182 APPENDIX I Taxonomic Support The following individuals generously supported this project with their time and expertise. Family Taxonomic Specialist Affiliated Institution Al lecul idae Serge Laplante Agr icul ture & Agr i food Canada , O t tawa Anob i idae Don Bright Agr icul ture & Agr i food Canada , Ot tawa Anthr ib idae Bob Sk idmore Agr icul ture & Agr i food Canada , Ot tawa Bostr ich idae Bob Sk idmore Agr icul ture & Agr i food Canada , Ot tawa Buprest idae Anthony Davies Agr icul ture & Agr i food Canada , Ot tawa Byrrh idae Laurent LeSage Agricul ture & Agr i food Canada , Ot tawa Bytur idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Canthar idae Ale S m e t a n a Agricul ture & Agr i food Canada , Ot tawa Carab idae Jeff Jarrod University of British Co lumbia , C a n a d a Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Cerambyc idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Cery lon idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Chrysomel idae Laurent LeSage Agr icul ture & Agr i food Canada , Ot tawa Ci idae Don Bright Agr icul ture & Agr i food Canada , Ot tawa C lamb idae Anthony Davis Agr icul ture & Agr i food Canada , Ot tawa Cler idae Bob Sk idmore Agr icul ture 8c Agr i food Canada , Ot tawa Coccine l l idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Colyd i idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Cory loph idae Anthony Davis Agr icul ture & Agr i food Canada , Ot tawa Cryp tophag idae Richard Leschen Landcare Research New Zea land Ltd. Cucu j idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Curcu l ion idae Bob Anderson Canadian M u s e u m of Nature, O t tawa Dermest idae Yves Bousquet Agr icul ture & Agr i food Canada , O t tawa Derodont idae Don Bright Agr icul ture & Agr i food C a n a d a , Ot tawa Dyt isc idae Ale S m e t a n a Agr icul ture & Agr i food Canada , Ot tawa Elater idae Ed Becker Agr icul ture & Agr i food Canada , Ot tawa Eroty l idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Eucnemidae Ed Becker Agr icul ture & Agr i food Canada , O t tawa Eucinet idae Don Bright Agr icul ture & Agr i food Canada , O t tawa Hister idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Hydraen idae Ale S m e t a n a Agricul ture & Agr i food Canada , Ot tawa Hydrophi l idae Ale S m e t a n a Agr icul ture & Agr i food Canada , Ot tawa Lampyr idae Ale S m e t a n a Agr icul ture & Agr i food Canada , O t tawa Lathr idi idae Fred And rews Cal i fornia Depar tment of Food and Agricul ture, Sacramento . Leiodidae Anthony Davis Agr icul ture & Agr i food Canada , Ot tawa Lucan idae Serge Laplante Agr icul ture 8c Agr i food Canada , Ot tawa Lycidae Ale S m e t a n a Agricul ture & Agr i food Canada , Ot tawa Melandry idae Darren Pol lock University of Mani toba, Winn ipeg Melyr idae Don Bright Agr icul ture &. Agr i food Canada , Ot tawa Mordel l idae Don Bright Agr icul ture &. Agr i food Canada , Ot tawa Myce tophag idae Yves Bousquet Agr icul ture &. Agr i food Canada , Ot tawa Family Taxonomic Special ist Affi l iated Institution Nemonych idae Don Bright Agr icul ture & Agr i food Canada , Ot tawa Nit idul idae Anthony Davies 183 O e d e m e r i n a e Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Phalacr idae Anthony Davies Agr icul ture & Agr i food Canada , Ot tawa Pse laph idae Anthony Davies Agr icul ture & Agr i food Canada , Ot tawa Pt inidae Anthony Davies Agr icul ture & Agr i food Canada , Ot tawa Pyrochro idae Ale S m e t a n a Agricul ture & Agr i food Canada , Ot tawa Pyth idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Rh izophag idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa Salp ing idae Darren Pol lock University of Mani toba, Winn ipeg Scaphid i idae Anthony Davies Agr icul ture & Agr i food Canada , O t tawa Scarabae idae Serge Laplante Agr icul ture & Agr i food Canada , O t tawa Scir t idae Ale S m e t a n a Agr icul ture & Agr i food Canada , Ot tawa Scoly t idae Don Bright Agr icul ture & Agr i food Canada , Ot tawa Scrapt i idae Darren Pol lock University of Mani toba, Winn ipeg S c y d m a e n i d a e Sean O'Keefe Moorhead State University, Kentucky Si lphidae Anthony Davies Agr icul ture & Agr i food Canada , O t tawa Sphaer i t idae Anthony Davies Agr icul ture & Agr i food Canada , Ot tawa Sph ind idae Ale S m e t a n a Agr icul ture & Agr i food Canada , Ot tawa Staphy l in idae Ale S m e t a n a Agr icul ture & Agr i food Canada , Ot tawa Anthony Davis Agr icul ture & Agr i food Canada , Ot tawa Stenot rache l idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Tenebr ion idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Darren Pol lock University of Mani toba, W i n n e p e g Te t ra tomidae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Throsc idae Serge Laplante Agr icul ture & Agr i food Canada , Ot tawa Trogoss i t idae Yves Bousquet Agr icul ture & Agr i food Canada , Ot tawa 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. 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. y=85.65x+27.8 ESTIMATOR OF THE SAMPLE RATIO r: r = Z X L Syi where Xi = beetle weight of the i sample yi = beetle number of the i t h sample ESTIMATED VARIANCE OF r: V( r) = N - n • _ 1 _ • U Nn L t x2 yi-iIXijL2 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 X j = 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: N - n • Sx 2 - rho s y • s 2 Nn x 2 y x ESTIMATION OF TOTAL BEETLE NUMBER FROM 1993 SAMPLES E(Y) = Rx E(Y) = 91.96 x V(y) = x 2 (V(r)) V(y) = x 2(4.37) Bound = + 1.96 (x 2 (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 o f 1 1 . 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 A P P E N D I X 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 A b u n d a n c e / Site harvest CN m L£5 Species - Increasing Abundance Family 0 i C L CO o Q_ to o Q_ to o Q_ to O CL Adalia bipunctata (L inneaus) Coccinel l idae 0 0 0.2 0. 0.3 Aleochara sekanai K l imaszewsk i Staphyl in idae 0 0.1 0 0. 0.2 Attica tombacina (Mannerhe im) Chrysomel idae 0 0 0 1.2 Amara lunicollis Schodte Carab idae 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 Ge rmar Elateridae 0 0 0.2 0. 0.2 Ampedus pullus Ge rmar Elateridae 0 3.2 3 5.5 Ampedus pullus Ge rmar Elateridae 0 . 1 2 1 6 Aphodius haemorrhoidalis (Linnaeus)/pectoral is LeConte Scarabae idae 0.1 0.1 0.2 0. 1 Bembidion grapii Gyl lenhal Carab idae 0 0 0 0. 0 . 4 Bolitopunctus muricatulus Staphyl in idae 0.9 1 2 1. 2 Bradycellus lecontei Csiki Carab idae 0 0 0 0 . 4 Bradycellus neglectus (LeConte) Carab idae 0 0.4 0.5 0. 0.7 Bradycellus nigrinus (Dejean) Carab idae 0.1 0.5 1.1 2. 1.8 Bradycellus nigrinus (Dejean) Carab idae 0 0 0 1. 5.8 Bryophacis smetanai Staphyl in idae 0 0.1 1 0. 1 Bryophacis smetanai Staphyl in idae 0 0 0 0. 0 . 4 Buperstis langi Mannerhe im Buprest idae 0 0 0 0. 0.3 Buperstis langi Mannerhe im Buprest idae 0 0 0 0 . 4 Buprestis nutialli Kirby Buprest idae 0 0.2 0.7 0. 1 Byturus unicolor Say Bytur idae 0 0 0.2 0. 0.3 Calitys scabra (Thunberg) Trogossi t idae 0.4 1.3 2 4. 4.3 Callidium cicatricosum Mannerhe im Cerambyc idae 0 0.1 0 0. 0.2 Callidium cicatricosum Mannerhe im Cerambyc idae 0 0 0 0. 0 . 2 Carphonotus testaceus Casey Curcul ionidae 0 0 0 0. 0 . 2 Cercyon sp#2 (cinctus) S m e t a n a Hydrophi l idae 0 0 0 0 . 4 Coccinella septumpunctata L inneaus Coccinel l idae 0 1 2.3 3. 2.3 Coccinella trifasciata perplexa Mulsant Coccinel l idae 0 0.1 0.5 0. 1.5 Colon asperatum Leiodidae 0 0 0 0. 0 . 6 Cododera sp.#514 Cerambyc idae 0 0 0.2 0. 0.2 Cosmosalia chrysocoma (Kirby) Cerambyc idae 0 0.1 0.1 0. 0.2 Crepidodera sp Chrysomel idae 0 0 0.2 0. 0.2 Cryptophagus sp#3 Cryptophag idae 0.3 0.3 1 1. 1.5 Ctenicera aeripennis (Kirby) Elateridae 0.2 3.5 20 24. 19.3 187 Ctenicera aeripennis (Kirby) Ctenicera angusticollis (Mannerhe im) Ctenicera kendalli Kirby Ctenicera nigricollis (Bland) Ctenicera pudica(V\I.J. B rown +propola columbiana)(Lecon\e) Ctenicera r. resplendens (Eschschol tz) Ctenicera r. resplendens (Eschschol tz) Ctenicera semimetallica (Walker) Ctenicera umbricola (Eschschol tz) Ctenicera umbricola (Eschschol tz) Cyphon concinnus (LeConte) Cyphon sp(p) Dalopius (nr) tristis W .J . B rown Dalopius (nr) tristis W .J . Brown Danosoma brevicorne (LeConte) Dicerca tenebrica (Kirby) Dichelonyx vicina (Fall) Didion punctatum (Melsheimer) Epuraea flavomaculata Makl in Eucnecosum tenue (LeConte) Gabrius picipennis (Makl in) Gyrophaena spp. Hadrobregmus americanus (Fall) Hemicoelus carinatus (Say) Heterothops conformis S m e t a n a Hydnobius pumilus LeConte Hydnobius sp# 1 Hydnobius sp# 1 Hydnobius sp#2 Hydnobius sp # 2 Laccobius borealis Cheary /earn D.C. Miller Laemophloeus biguttatus (Say) Lelinohesperus borealis Leoides collaris (LeConte) Leoides collaris (LeConte) Leoides puncticollis/curvata C.G. T h o m s o n / M a n n e r h e i m Leoides sp # 3 Leptusa sp Limonius pectoralis LeConte Lordithon cascadensis (Maikin) Lordithon t. thoracicus (Fabric ius) Megasemum asperum (LeConte) Neoclytus m. muricatulus (Kirby) Pediacus depressus (Herbst) Phausis rhombica Fender Platycerus marginalis Casey Ptinus californicus Pic Pygoleptura n. nigrella (Say) Quedius m. molochinoides Smetana Elateridae 0 Elateridae 0.5 Elater idae 0 Elateridae 1.5 Elateridae 5.5 Elateridae 0.2 Elateridae 0 Elateridae 0 Elateridae 0.7 Elateridae 0 . 4 Scirt idae 0 Scirt idae 0 Elateridae 0 Elater idae 0 Elater idae 0 Buprest idae 0 Scarabaeidae 0 Coccinel l idae 0 Nitidulidae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Anobi idae 0 Anobi idae 0 Staphyl in idae 0 Leiodidae 0 Leiodidae 0 Leiodidae 0 Leiodidae 0 Leiodidae 0 Hydrophi l idae 0 Cucuj idae 0 Staphyl in idae 0 Leiodidae 0 Leiodidae 0 Leiodidae 0 Leiodidae 0 Staphyl in idae 0 Elateridae 0 Staphyl in idae 0 Staphyl in idae 0 Cerambyc idae 0.1 Cerambyc idae 0 Cucuj idae 0 Lampyr idae 0 Lucanidae 0 Ptinidae 0 Cerambyc idae 0 Staphyl in idae 0 6 12 4. 1 4 . 4 0 .9 0 .6 1.2 0 0 0. 0 . 4 2 .5 3.7 3. 6.7 9 .3 10.4 2 7 . 4 5 . 7 3.8 1 1 . 6 18. 15.8 4 4 1 3 . 8 0.2 0 .4 0. 0.7 11.6 18 .8 14. 5 0 . 8 4 6 2 4 . 3 3 . 8 0 0 0 . 4 0 0 2. 1 0.5 1.5 1.7 0 0 1. 2 . 6 2 0 3. 5 . 6 1 3 1 8 . 8 0 0 0.5 0 0 0 . 4 0 0 0. 0 . 4 0.1 0 .2 0. 0.2 0 0 0. 1 0 0 0. 0 . 4 0 0 0. 1.4 0.1 0.1 0. 0.2 0 0 0. 0 . 2 0 1 0. 1 0.5 0 .7 0. 1.5 0 0 0. 1 0.2 0 .3 0. 0.7 0 0 0. 0.8 0 0 0. 0.4 0 0 0. 1.5 0 0 0 . 4 0.1 0 0. 0.2 0 0 0. 0.4 0 .2 0.3 0. 0.5 0 0.1 0. 0.2 0 0 0. 0 . 6 0 0 0. 0 . 2 0 0 0. 0.3 0.1 0 .2 0. 0.7 0 .3 0 .8 0. 0.8 0.1 0.1 0. 0.3 0 0 0. 0 . 2 0 0 0. 0.2 0 0 0.8 0 0 0. 0 . 2 0 0 0. 0 . 8 0 1 2. 1 188 Quedius rusticus/vilis S m e t a n a Scaphisoma castaneum Motschulsky Scolytus piceae (Swaine) Semijulistus ater (LeConte) Sericus brunneus sp#611 Stenichnus californicus Motschulsky Stenichnus californicus Motschulsky Strictoleptura canadensis cribripennis (LeConteJ Syntomus americanus (Dejean) Tachinus thruppi Hatch Tachyta rana (Casey/Say) Teretruis montanus Horn Tomoxia borealis (LeConte) Tribolium audax Halstead Trichodes ornatus hartwegianus A. Whi te Trixagus sp Trixagus sp (carnicollis (Schaef fer ) j Tropideres fasciatus (Olivier) Staphyl in idae 0 0 0 0. 0 . 2 Scaphidi idae 0 0 0 0. 0 . 2 Scolyt idae 0 0 0 0. 0 . 2 Melyr idae 0 0 2 2. 2 Elateridae 0 0 0.1 0. 1.3 Scarabae idae 0 0 0 0. 0 . 2 Scydmaen idae 0 0.2 0 .2 0. 0.3 Scydmaen idae 0 0 0 0 . 4 Cerambyc idae 0.1 3 .6 6 .6 13 . 8 .3 Carab idae 0 0 0 0. 0 . 2 Staphyl in idae 0 0 0 0. 0 . 2 Carab idae 0 0 0 0. 0.2 Hister idae 0 0.2 0.4 1 . 2.2 Mordel l idae 0 0 0 0. 0 . 2 Tenebr ion idae 0 0 1 0. 1.6 Cleridae 0 0 0 0 . 4 Throsc idae 0 0 0 0 . 4 Throsc idae 0 0.1 0 0. 0.2 Anthr ib idae 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 harves T— CM CO in Species - Decreasing Abundance Family i— Q. to o Q_ OO O Q_ W o CL to o 0. Abstrulia (nr) veriegatta Casey Tet ra tomidae 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 Scrapt i idae 25.6 8.7 9.6 3.2 3.2 Aphodius leopardus Horn Scarabaeidae 0 . 7 0 0 0 0 Athous nigropilis Motschulsky Elater idae 0 . 3 0 0 0 0 Atrecus macrocephalus (Nordmann) Staphyl in idae 0.6 0.3 0.3 0.2 0 Caenocara scymnoides LeConte Anobi idae 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 Mannerhe im Cucuj idae 4 4 1.3 0.4 1 Dicentrus bluthneri LeConte Cerambyc idae 0.2 0.2 0.1 0 0 Dicentrus bluthneri LeConte Cerambyc idae 0 . 3 0 0 0 0 Dienopteroloma subcostatum (Makl in) Staphyl in idae 0 . 3 0 0 0 0 Drasterius debilis LeConte Elateridae 3 4 1 0 . 6 0 . 4 Dryocetes caryi/schelti Hopk ins /Swaine Scolyt idae 0.2 0.2 0 0 0 Earota sp. Staphyl in idae 1 1 0 0 0 Earota sp. Staphyl in idae 0 . 3 0 0 0 0 Eleates explanatus Casey Tenebr ion idae 0.3 0.2 0 0 0 Emmesa stacesmithi Hatch Melandry idae 0.4 0.1 0.1 0 0 Enicmus mendax Lathridi idae 0.1 0.1 0 0 0 Enicmus mendax Lathridi idae 0.3 0 0 0 0 Enicmus tenuicornis LeConte Lathridi idae 2.6 2.5 2.1 1.2 1.3 Eupraea terminalis Mannerhe im Nit idulidae 0.2 0 0 0 0 Eusphalerum spp (mostly pothos (Mannerhe im)) Staphyl in idae 202 2.9 2.7 1.1 3.7 Hallomenus sp Scrapt i idae 0 . 3 0 0 0 0 Lathridius n sp Lathridi idae 0.5 0.2 0 0 0 Lathridius n sp Lathridi idae 0 . 3 0 0 0 0 Lordithon fungicola Campbel l Staphyl in idae 1.2 0.4 0.2 0 0 Omalium spit 1 Staphyl in idae 0.1 0.1 0 0 0 Omosita discoidea (Fabr ic ius) Nit idulidae 3.5 1.3 0.6 0.1 0 Odhocis punctatus Casey Ci idae 1.4 0.4 0.5 0.3 0.2 Pelecomalium testaceum (Mannerhe im) Staphyl in idae 4.5 1.8 0.6 0.2 0.5 Phryganophilus collaris LeConte Melandry idae 0.1 0.1 0 0 0 Pidonia scripta (LeConte) Cerambyc idae 0.5 0 0 0 0 Placusa tacomae Staphyl in idae 0.1 0.1 0.1 0 0 Pytho sp#2 Pythidae 0.3 0.1 0.1 0 0 Quedius criddlei (Casey) Staphyl in idae 0.4 0 0 0 0 Quedius plagiatus Mannerhe im Staphyl in idae 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 Mannerhe im Rhizophagidae 2.6 0.5 0.1 0.2 0.2 190 Salebius nr minax Scierus annectans LeConte s p # 10 Spondylis upiformis Mannerhe im Stephostethus breviclavus (Fall) Stephostethus liratus (LeConte) Syneta pilosa W . J . Brown Tachinus frigidus Er ichson Tetratoma concolor LeConte Tetropium velutinum LeConte Thanasimus undatulus (Say) Thymalus marginicollis Chevrolet Thymalus marginicollis Chevrolat Trachysida a. aspera (LeConte) Trypodendron retusum (LeConte) Trypodendron retusum (LeConte) Trypodendron rufitarsis (Kirby) Trypodendron rufitarsis (Kirby) Xylechinus montanus B lackman Xylechinus montanus B lackman Cryptophagidae 0 . 6 Scolyt idae 14.9 Nitidulidae 0.1 Cerambyc idae 4.3 Lathridi idae 0.4 Lathridi idae 0.9 Chrysomel idae 0 . 3 Staphyl in idae 0.2 Tetratomidae 1.9 Cerambyc idae 1.3 Cler idae 120 Trogossi t idae 2.7 Trogossi t idae 4 . 3 Cerambyc idae 0.3 Scolyt idae 2 Scolyt idae 1.3 Scolyt idae 0.7 Scolyt idae 0 . 3 Scolyt idae 5.6 Scolyt idae 0 . 7 0 0 0 0 9.6 4.3 0.6 0.8 0.1 0.1 0 0 3.1 0.7 1 1.7 0.4 0.2 0.3 0 0.5 0.2 0.5 0.2 0 0 0 0 0.1 0.1 0 0 0.2 0.3 0 0 2 1 0 0 116 75.8 29.4 28.5 0.6 0.2 0 0 4 2 0 0 0.2 0.3 0 0 0.9 0.1 0.1 0.2 1 0 0 0 0 0.1 0 0 0 0 0 0 0.2 0.1 0 0 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 A b u n d a n c e / Site Species with increasing then decreasing abundance sp # 328 Abstrulia (nr) veriegatta Casey Acidota crenata (Fabric ius) Acmaeops p. proteus (Kirby) Agathidium depressum Fall /obtusum Hatch Agathidium difformis (LeConte) Agathidium difformis (LeConte) Agriotella occidentalis W . J . B rown Aleochara gracilicornis Bernhauer Aleochara rubricalis Casey Aleochara suffosa (Casey) Aleocharinae (misc.spp) Allandrus populi Pierce Altica tombacina (Mannerhe im) Amara discors Kirby Amara erratica (Dutschmid) Amara familiaris (Dutschmid) Amara laevipennis Kirby Amara latior (Kirby) Amara littoralis Mannerhe im Amara littoralis Mannerhe im Amara lunicollis Schodte Amischa sp. Ampedus (nr) moerens (LeConte) Ampedus behrensi + phelpsi (Horn) Ampedus brevis (Van Dyke) Ampedus brevis (Van Dyke) Ampedus mixtus (miniipennis?) (Herbst) Ampedus moerens (LeConte) Ampedus nigrinus (Herbst) Ampedus nigrinus (Herbst) Ampedus occidentalis Lane Ampedus occidentalis Lane Anaspis sp#2 Anisotoma globososa Hatch Anotylus rugosus (Farbr ic ius) Anthaxia inomata (Randal l ) Antherophagus sp#1 Antherophagus sp#2 Antherophagus sp#2 £ i n CD £Z CN CO ?F Family CD u_ Q_ t o o CL W O CL t o o CL t o o CL Scarabae idae 0 . 1 2 1 0 0 Tetra tomidae 1.6 3 0 0 0 Staphyl in idae 0.2 0.6 1.9 2.3 0.3 Cerambyc idae 0 0.4 0.5 0.2 0.3 Leiodidae 2.7 2 6 1 1.2 Leiodidae 0 0 0.2 0.1 0 Leiodidae 0.3 1 0 0 0 Elater idae 0 0 0 0 . 4 0.2 Staphyl in idae 0 0.3 0.2 0.2 0.2 Staphyl in idae 0 0.1 0.5 0.1 0 Staphyl in idae 0 0.9 0.4 0.1 0.2 Staphyl in idae 1 2.6 0.9 0.1 0 Anthr ib idae 0 0.1 0.1 0.1 0 Chrysomel idae 0 0.2 0.6 0.3 0.2 Carab idae 0 0 0.1 0.1 0 Carab idae 0 1 1 0 . 6 0.6 Carab idae 0 0.1 0.1 0.1 0 Carab idae 0 0.4 0.6 0 0 Carab idae 0 0 0.2 0 0 Carab idae 0 0.3 0.5 1.2 0.3 Carab idae 0 0 0 2 . 2 0 Carab idae 0 0.5 0.6 0.8 0.5 Staphyl in idae 0 0 0.2 0.3 0 Elateridae 0 4 8.4 1.7 3.7 Elateridae 0 2.9 14.5 4.2 7.2 Elateridae 0.7 3.5 5.5 4.3 4 Elateridae 2 . 4 6 5 3 . 4 3.6 Elateridae 0 0.9 1.6 0.8 1.2 Elateridae 0 1.8 1.2 1.1 1.2 Elateridae 0.6 14.5 23.1 14 10.3 Elateridae 0 . 1 30 20 2.8 1 2 . 8 Elater idae 0.2 3.5 5.6 3.7 3.8 Elater idae 0 25 2 1 4 . 6 4 . 8 Scrapt i idae 0 1.1 1.2 1.4 0.5 Leiodidae 0 0.5 1.2 1.2 0.2 Staphyl in idae 0 0.5 0.2 0.4 0 Buprest idae 0.2 1.7 0.5 0.3 0 Cryptophag idae 0 4 0 0 0 Cryptophag idae 0.1 0.5 0.8 0.1 0.2 Cryptophag idae 0 0 0 0 . 4 0 192 Asemum striatum (L inneaus) Asemum striatum (L inneaus) Atheta dentata Athous rufiventris rufiventris (Eschschol tz) Athous rufiventris rufiventris (Eschschol tz) Atomaria sp #10 Atomaria spit 12 Atomaria sp # 12 Atomaria sp#6 Bembidion grapii Gyl lenhal Bembidion nigripes (Kirby) Bembidion tetracolum Say Bembidion versicolor (LeConte) Bhyrrhus sp Bisnius picicornis Bius estriatus (LeConte) Bradycellus congener (LeConte) Bryophacis arcticus Bryophacis punctulatus Bryophacis punctulatus Bryophacis spp Buprestis lyrata Casey Caenocelis sp# 1 Calitys scabra (Thunberg) Calopus angustus LeConte Calyptomerus oblongulus Mannerhe im Canifa sp# 1 Carphacis nepigonensis (Bernhauer) Carphacis nepigonensis (Bernhauer) Carphoborus vandykei Bruck Cephaloon tenuicorne LeConte Cerycon sp#1 (herceus frigidus) Smetana Cerylon castaneum Say Cerylon castaneum Say Ceutorhynchus punctiger Gyl lenhal Chrysobothris carinipennis LeConte Chrysobothris carinipennis LeConte Cis angustus Hatch Colon asperatum Colon magnicolle Mannerhe im Corticaria n sp Corticeus praetermissus (Fall) Corticeus subopacus (Wall is) Corticeus subopacus (Wall is) Corticeus tenuis (LeConte) Cortodera m. militaris (LeConte) Cortodera sp. #514 Cosmosalia chrysocoma (Kirby) Cossonus pacificus V a n Dyke Cryphalus ruficollis Hopkins Cerambyc idae 0.1 Cerambyc idae 0 . 1 Staphyl in idae 0.3 Elateridae 0.9 Elateridae 2 . 1 Cryptophag idae 0 Cryptophagidae 0.1 Cryptophagidae 0 Cryptophagidae 0 Carabidae 0 Carabidae 0 Carabidae 0 Carab idae 0 Byrrhidae 0 Staphyl in idae 0.3 Tenebr ion idae 0 Carab idae 0 Staphyl in idae 0 Staphyl in idae 0.1 Staphyl in idae 0 Staphyl in idae 0 Buprest idae 0 Cryptophagidae 0.3 Trogossi t idae 0 . 4 Oedemer inae 1 Clambidae 0 Melandry idae 0 Staphyl in idae 0.4 Staphyl in idae 0 . 1 Scolyt idae 0 Cephalo idae 0.3 Hydrophi l idae 0 Cerylonidae 0.6 Cery lonidae 0 . 9 Curcul ionidae 0 Buprest idae 0 Buprest idae 0 Ciidae 0 Leiodidae 0 Leiodidae 1.2 Lathridi idae 4.3 Tenebr ion idae 0.1 Tenebr ion idae 0.2 Tenebr ion idae 0 . 1 Tenebr ion idae 0 Cerambyc idae 0.4 Cerambyc idae 0 Cerambyc idae 0 Curcul ionidae 0.2 Scolyt idae 0.1 5.1 0.3 0 0 8 0 0 0 2.4 1.4 0.4 0 1.5 7.1 1 1.8 4 3 2.4 2.6 0 0 0.3 0 0.4 0.1 0.1 0 0 1 0 . 2 0 0.1 0.1 0.1 0 0.8 0.6 0.4 0 0 0.1 0.1 0 0.1 0.4 0 0 0 0.3 0.2 0 0.1 0.1 0 0 1.7 2 0 0 1.5 0.3 0 0 0 0.2 0 0 0.2 0 0 0 0.3 0.4 0.1 0 1 1 0 . 2 0 0.1 0.2 0.2 0 0 11 8 . 4 7.2 2.3 1.6 0.6 0.5 0 5 1 1.8 1.5 3 1 0.5 0.1 0.1 0 0 0 0.2 0.3 0 1.2 1.5 0.3 0.5 1 2 0 0 . 4 0.5 0.3 0.3 0 1.2 1.9 1.1 0.7 0.2 0.1 0.6 0 0.8 1.3 0.9 0.7 2 2 0 . 8 0 . 4 0.1 0.1 0 0 0 0.3 0.4 0 0 0 0 . 6 0 0.2 0.6 0.2 0.3 0.4 0.4 0.4 0.3 1.4 1.5 0.6 0.7 5.4 5.5 1.9 0.5 0.3 0.2 0 0 0.2 0.4 0.1 0 2 0 0 0 0 0 0 . 4 0 1.5 1.5 0.3 0.2 0 5 0 . 4 0 . 2 0 0 0 . 4 0 1.1 0.6 0.5 0.2 0.7 0.1 0.2 0 193 Cryptophagus sp#4 Ctenicera bombycina (Germar ) Ctenicera lobata (Eschschol tz ) Ctenicera lobata (Eschschol tz ) Ctenicera lutescens (Fa\\)/sagitticollis (Eschschol tz) Ctenicera nebraskensis (Bland) Ctenicera nitidula (LeConte) Ctenicera triundulata (Randal l ) Ctenicera volitans (Eschschol tz) Cucujus claviceps Mannerhe im Cylistus coarctatus (LeConte) Cylistus coarctatus (LeConte) Cyphon sp(p) Cytursa luggeri Danosoma brevicorne (LeConte) Dendroctonus pseudotsugae Hopkins Dendroctonus pseudotsugae Hopkins Dendroctonus rufipennis (Kirby) Dendrophagus cygnaei Mannerhe im Dermestes talpinus Mannerhe im Dicerca tenebrica (Kirby) Dicerca tenebrosa (Kirby) Didion punctatum (Melsheimer) Dienopteroloma subcostatum (Makl in) Dolichocis manitoba Dury Drasterius debilis LeConte Dryocetes affaber (Mannerhe im) Dryocetes affaber (Mannerhe im) Dryocetes autographus (Ratzeburg) Dryocetes autographus (Ratzeburg) Dryocetes confusus Swa ine Dryocetes confusus Swa ine Dyctyopterus spp Elaphrus americanus Dejean Ellychnia corrusca ( L i n n e a u s ) Ellychnia corrusca (L inneaus) Enoclerus nr. Scheaferi Barr Enoclerus nr. Scheaferi Barr Enoclerus sphegeus (Fabr icus) Epiphanis sp # 338 Epuraea planulata Er ichson Epuraea planulata Er ichson Epuraea spit 1 Epuraea sp# 1 Eupraea truncatella Mannerhe im Eupuraea spit 2 Evodinus monticola vancouveri Casey Gabrius picipennis (Makl in) Glischrochilus confluentus (Say) Glischrochilus quadrisignatus (Say) Cryptophagidae 2 . 1 1 2 0.6 0.8 Elateridae 0 0.3 0.4 0.6 0 Elateridae 0 0.1 0.1 0 0 Elateridae 0 0 0 0 . 4 0 Elateridae 0.2 1.5 0.6 0.5 0.2 Elateridae 0.4 5.6 8.8 9.6 5.2 Elateridae 0 0.1 0.4 0.3 0 Elateridae 0 0 0.1 0.1 0 Elateridae 0 . 4 4 1 2.2 0.8 Cucuj idae 1.9 3 6 0.2 0 Hister idae 0 0.6 0.7 0 0 Hister idae 0 1 1 0 0 Scirt idae 0.1 0.3 1.1 1.5 0.7 Leiodidae 0 0 0 0 . 4 0 . 2 Elateridae 0 1.7 3.3 5.6 3.8 Scolyt idae 7699 12859 10362 4152 1137 Scolyt idae 6 . 4 1 6 15 7 . 6 2 Scolyt idae 0 0.1 0.2 0.1 0 Cucuj idae 1.7 5 1.5 0.5 0.3 Dermest idae 0.1 0.2 0.2 0 0 Buprest idae 0 2.2 9.4 9.6 7.5 Buprest idae 0 3.3 3 0.9 1 Coccinel l idae 0 0.1 0.3 0.2 0.2 Staphyl in idae 0.1 0.5 0.1 0 0 Ciidae 0.1 0.7 0.4 0.2 0.2 Elateridae 0.5 1.1 0.2 0.5 0 Scolyt idae 0.5 7.4 1.4 0.8 0.2 Scolyt idae 0.6 6 4 0.8 0 . 4 Scolyt idae 1.1 18.8 5.9 2.1 0.7 Scolyt idae 0.6 1 4 4 2 2 . 4 Scolyt idae 0.3 4.1 0.3 0 0 Scolyt idae 0 . 1 1 0 0 0 Lycidae 1 2 6 0 . 4 0 . 8 Carab idae 0 0 0.1 0.1 0 Lampyr idae 0 0 0.3 0 0 Lampyr idae 0 0 1 0.2 0 Cler idae 0.1 0.4 0 0 0 Cler idae 0 3 0 0 0 Cleridae 0.2 1 0.5 0 0 Eucnemidae 0 0.1 0.1 0.1 0 Nitidul idae 0.1 1.5 1.5 0.3 0.2 Nitidul idae 0 . 3 3 3 0.2 0 . 4 Nitidul idae 0.2 3.1 0.9 0.3 0.2 Nitidul idae 0 . 3 2 0 0.8 0 . 2 Nitidulidae 0.3 2.1 1.4 0.1 0.2 Nitidulidae 0 0 0.1 0.1 0 Cerambyc idae 0 . 9 3 1 0 0 Staphyl in idae 0 2.4 5.8 2.4 1.2 Nitidul idae 0.1 2.6 0.2 0.1 0.2 Nitidul idae 0 0.4 0.5 0 0 194 Gnathacmaeops pratensis (Laichart ing) Gnathoncus barbatus Bosquet & Laplante Gnathoncus communis (Marseul ) Gnathotrichus retusus LeConte Gnathotrichus retusus LeConte Grypeta sp. Gyrophaena spp. Hadrobregmus americanus (Fall) Hadrobregmus quadrulus (LeConte) Hallomenus sp Hapalaraea dropephylla Hapalaraea megadhroides (Fauvel ) Hapalaraea megadhroides (Fauvel ) Hapalaraea sp #1 Harpalus animosus Casey Harpalus laevipes Harpalus somnulentus Dejean Harpalus somnulentus Dejean Helophorus orientalis Motscho lsky /sempervar ians A n g u s Henoticus sp# 1 Heterothops conformis S m e t a n a Heterothops sp Hoppingiana sp (hudsonica) (LeConte) Hoppingiana sp (hudsonica) (LeConte) Hydnobius pumilus LeConte Hydraena sp (pacifica) Perk ins Hydrobius fuscipes (L inneaus) Hydrobius fuscipes (L inneaus) Hydroporus sp# 1 Hydroporus sp#3 Hylastes nigrinus (Mannerhe im) Hylastes nigrinus (Mannerhe im) Hylastes ruber Swa ine Hylurgops porosus (LeConte) Hylurgops porosus (LeConte) Hylurgops rugipennis Mannerhe im/F i tch Hypnoidus bicolor (Eschschol tz) Ips perturbatus (Eichhoff) Ips pedurbatus (Eichhoff) Ips pini (Say) Ips tridens (Mannerhe im) Judolia m. montivagens (Couper) Judolia m. montivagens (Couper) Laccobius borealis Cheary /card D.C. Miller Lathridius ventralis Lathrobium negrum LeConte Leiodes strigata (LeConte) Lelinohesperus borealis Leoides puncticollis C.G. T h o m s o n /curvata Mannerhe im Cerambyc idae 0 0.3 0 .4 0.1 0 Hister idae 0 0.5 0 .2 0 0 Hister idae 0 0 0.1 0.1 0 Scolyt idae 2.2 1 5 . 7 3 0 . 7 12.7 0.7 Scolyt idae 0 . 6 4 2 4 . 4 1.4 Staphyl in idae 0 0.2 0 0 0 Staphyl in idae 0 0.5 0 .2 0 .2 0 Anobi idae 0 0.3 0 .5 1.1 0 Anobi idae 0 0 0.2 0.1 0 Melandry idae 0.1 0.1 0 .3 0 0 Staphyl in idae 0 0.2 0 0 0 Staphyl in idae 0.1 1 0 0.1 0 Staphyl in idae 0 . 4 2 0 0 0 Staphyl in idae 0.1 0.2 0 0 0 Carab idae 0 0 0 1 0 Carab idae 0 0.1 0 .5 0 0 Carab idae 0 0.1 0 .5 0.2 0 Carab idae 0 0 0 0 . 6 0 . 2 Hydrophi l idae 0 0.4 0.4 0.2 0.3 Cryptophagidae 0.1 0 .9 0.4 0 0 Staphyl in idae 0 3.2 5 .8 1.5 0.2 Staphyl in idae 0 0 0.3 0 0 Melyr idae 0.2 0 .2 1.1 0 .3 0 Melyr idae 0 0 0 0 . 4 0 . 2 Leiodidae 0 0.5 0 .9 0 .7 0.3 Hydraenidae 0 0 0.2 0.2 0 Hydrophi l idae 0 0.3 0.3 0.2 0 Hydrophi l idae 0 0 0 0 . 6 0 . 2 Dytiscidae 0 0 0.1 0.1 0 Dytiscidae 0 0.1 0 .2 0.1 0 Scolyt idae 13 .2 1 4 0 1 2 0 4 4 . 2 8.2 Scolyt idae 6 . 1 2 2 4 1 3 1 1 . 4 1 3 . 2 Scolyt idae 5.1 5 4 . 2 3 4 . 5 9 10.3 Scolyt idae 0 1 0 5 2 0 . 4 3.5 1.7 Scolyt idae 0 . 1 82 2 0 . 2 0 . 6 Scolyt idae 0 . 1 1 0 0 0 Elateridae 0 0 1 1.6 0 . 2 Scolyt idae 0.1 5 .3 0 .8 0 0.2 Scolyt idae 0 4 2 0 . 6 0 . 2 Scolyt idae 0.1 1.7 0 .5 0 0.5 Scolyt idae 0 3.4 0 .9 0.1 0 .3 Cerambyc idae 0 0.4 2 .5 0.7 0 .3 Cerambyc idae 0 . 1 1 10 0 . 4 0 . 6 Hydrophi l idae 0 0 0 0.4 0 Lathridi idae 0 0.2 0.1 0 0 Staphyl in idae 0 0.3 0 0 0 Leiodidae 0 4 . 8 1.4 0 .5 0.7 Staphyl in idae 0 0.2 0.1 0 .2 0 Leiodidae 0 0 1 0 . 6 0 . 2 195 Leoides rufipes (Gebler) Leoides rufipes (Gebler) Leptusa sp Limonius pectoralis LeConte Medon sp. (nr. Pallescens) Megadhrus angulicollis Megasemum asperum (LeConte) Megatoma sp (cylindrica) (Kirby) Megatoma sp (cylindrica) (Kirby) Megatoma verigatta (Horn) Melandrya striata Say Melanophila drummondi (Kirby) Microbregma e. emarginatum (Duf tschmid) Microbregma e. emarginatum (Duf tschmid) Micropeplus smetanai Campbel l Molamba obesa Casey Monochamus spp Mycetophagus distinctus Hatch Mycetoporus brunneus (Marsham) Mycetoporus rufohumoralis Myrmecocephalus arizonicus (Casey) Myrmecocephalus arizonicus (Casey) Myrmedophila americana (LeConte) Neanthophlax mirificus (Bland) Negastrius tumescens LeConte Negastrius tumescens LeConte Neoclytus m. muricatulus (Kirby) Nitidotachinus tachyporus Nudobius cephalus (Say) Odontosphindus clavicornis Casey Olophrum boreale (Paykul l ) Olophrum boreale (Paykul l ) Olophrum consimile (Gyl lenhal) Omalium sp # 2 Omalium spp. Omosita discoidea (Fabric ius) Orchesia (nr) castanea Orsodacne atra (Ahrens) Odhotomicus caelatus (Eichhoff) Odhotomicus caelatus (Eichhoff) Ostoma ferrugina (Linneaus) Oxytelus sp. Pachyta lamed liturata Kirby Pactopus hornii (LeConte) Pactopus hornii (LeConte) Paromalus mancus Casey Pediacus depressus (Herbst) Pediacus fuscus Er ichson Pediacus fuscus Er ichson Pelecomalium testaceum (Mannerhe im) Leiodidae 0 Leiodidae 0 . 1 Staphyl in idae 0.1 Elateridae 0 Staphyl in idae 0 Staphyl in idae 0.2 Cerambyc idae 0 . 6 Dermest idae 0.6 Dermest idae 0 . 9 Dermest idae 1.1 Melandry idae 0.1 Buprest idae 0 Anobi idae 0.6 Anobi idae 0 . 7 Staphyl in idae 0 Cory lophidae 0 Cerambyc idae 0 Mycetophag idae 0 Staphyl in idae 0 Staphyl in idae 0.4 Staphyl in idae 0 Staphyl in idae 0 Cryptophag idae 0 Cerambyc idae 1.7 Elateridae 1 Elater idae 0 Cerambyc idae 0 Staphyl in idae 0 Staphyl in idae 0 Sphindidae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Nitidulidae 0 . 3 Scrapt i idae 0.1 Chrysomel idae 0.1 Scolyt idae 0.1 Scolyt idae 0 Trogossi t idae 0 . 3 Staphyl in idae 0 Cerambyc idae 0 Throsc idae 0.4 Throsc idae 0 . 1 Histeridae 0.6 Cucuj idae 0 Cucuj idae 0.2 Cucuj idae 0 Staphyl in idae 0 . 3 0 0.2 0 .2 0 1 1 0 0.8 0 .2 0 .2 0.1 0 0 0.2 0.2 0 0.1 0 .4 0.1 0.3 1.4 0 .4 0.2 0 3 1 0 . 4 0 . 8 3.7 0 .9 0 .7 0 2 3 0 0 2.1 1.8 0 .8 0.2 0.1 0 .4 0.2 0.3 0 .5 0 .4 0.2 0 2.3 2 . 4 0.7 0 1 6 0 . 4 0 . 2 0.5 0.1 0.2 0 0.2 0 0 0 1.4 0 .9 0.3 0.3 0 .6 0 .5 0.3 0.3 0 0.3 0 0 1 0.5 0.4 0.5 0.1 0.1 0.1 0 0 0 0 . 4 0 0.4 0 .5 0.2 0.3 1 7 3 4 . 6 5 2 9 . 1 1 0 . 6 3.4 5 .8 5 6 10 1 3 . 6 0 0 0 . 6 0 . 2 0 0.1 0.1 0 3.1 1.8 0 .9 0 .3 0 .3 1.2 0 .5 0 0.1 0.1 0 .2 0 0 0 0 . 4 0 0 1 0 . 6 0 . 6 0.1 0.1 0 0 0.3 0 0 0 2 0 0 0 0.2 0.1 0 .2 0 0.7 0 .2 0 .5 0 .3 4 5.7 1.5 0.2 10 2 0 .6 0 . 2 1 1 0 . 4 0 . 4 0.7 0 0 0 1.9 0.4 0 0 0.6 1.1 0 .3 0 .7 2 2 0 . 2 1 4 .3 0 .9 0.1 0 .2 0 .3 0.4 0 .2 0.2 4 . 6 9 .2 1.4 1.5 4 4 1.8 0 . 8 1 5 2 0 . 4 0 . 2 196 Phaleromela verigata Tr ip lehorn Philodrepa (?) Dropephylla sp (nrlongula) Philonthinii spp Philonthus furvus N o r d m a n n Phloeophagus canadensis V a n Dyke Phloeotribus lecontei/picea Sched l /Swaine Phyllotreta striolata (Fabr icus) Pissodes (nr) fiskei Hopkins Pissodes fasciatus LeConte Pissodes striatulus (Fabr icus) Pissodes striatulus (Fabr icus) Pissodes striatulus dubius Randal l Pityogenes hopkinsi Swa ine Pityogenes plagiatus (LeConte) Pityokeines minutus (Swaine) Pityokteines elegans Swa ine Pityophthorus nitidulus S w a i n e ^ tuberculatus EschhoffJ Pityophthorus nitidulus S w a i n e ^ tuberculatus Eschhoff) Pityophthorus opaculus LeConte Pityophthorus opaculus LeConte Pityophthorus pseudotsugae Swaine Pityopthorus aquilus B lackman (+ aplanatus) Platycerus marginalis Casey Platysoma coarctatum LeConte Platysoma leconti Marseul Plesiocis sp Podabrus piniphilus (Eschschol tz) Pogonocherus penicillatus LeConte Pogonocherus penicillatus LeConte Poliaenus oregonus (LeConte) Polygraphus convexifrons W o o d Polygraphus rufipennis (Kirby) Polygraphus rufipennis (Kirby) Priognathus monilicornis LeConte Pseudohadrotoma sp (perversa) (Fall) Pseudohylesinus nebulosus LeConte Psyllobora vigintimaculata (Say) Pterostichus adstrictus Eschschol tz Pytho sp # 1 Pytho sp # 1 Pytho spit 2 Quediini spp Quedius disticalios Quedius m. molochinoides Smetana Quedius rusticus/vilis S m e t a n a Quedius transparens Motschulsky Quedius velox S m e t a n a Quedius velox S m e t a n a Rhagium inquisitor (L inneaus) Tenebr ion idae 0 Staphyl in idae 0.1 Staphyl in idae 0.3 Staphyl in idae 0 Curcul ionidae 0 Scolyt idae 0 Chrysomel idae 0 Curcul ionidae 0 Curcul ionidae 0.2 Curcul ionidae 0 Curcul ionidae 0 . 1 Curcul ionidae 0 Scolyt idae 0.1 Scolyt idae 0.1 Scolyt idae 0 Scolyt idae 0 Scolyt idae 0.1 Scolyt idae 0 . 3 Scolyt idae 0 Scolyt idae 0 Scolyt idae 0.1 Scolyt idae 0 Lucanidae 0 Hister idae 0 Hister idae 0 Ciidae 0 Canthar idae 0.1 Cerambyc idae 0 Cerambyc idae 0 Cerambyc idae 0 Scolyt idae 0.4 Scolyt idae 3.2 Scolyt idae 0 . 3 Pythidae 0 Dermest idae 0 Scolyt idae 4.3 Coccinel l idae 0 Carab idae 0 Pythidae 0 Pythidae 0 Pythidae 0 . 1 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0.3 Staphyl in idae 0 Staphyl in idae 4.4 Staphyl in idae 4 . 7 Cerambyc idae 0 0.1 0.1 0 0 1.1 6.9 4.6 4.2 1.2 2 0.7 0.3 0 0.1 0.1 0 0 0.2 0 0 1.5 0.3 0.2 0 0.1 0.1 0 0 0.2 0 0 0 1.6 0.5 0 0 1.3 0.3 0 0 7 0 0 0 0.5 0 0 0 0.1 0.4 0.1 0.2 4.2 1 0.3 0 0.5 0.5 0.5 0 0 0.3 0 0 0.8 2.2 0.9 0.5 2 1 1.4 0 . 4 0.6 0.3 0.4 0.2 0 0 0 . 6 0 3.2 0.5 0.2 0 2 0 0 0 0.1 0.7 0 0 0.2 0.1 0 0 0.1 0.5 0.5 0 0.3 0.1 0 0 0.4 0.5 0.1 0.3 1.5 0.3 0 0 11 0 0 0 1 1 0 0 1 0.2 0.1 0 43.2 21.1 7.8 4.3 12 6 5 . 8 2 . 2 1.1 0.3 0.1 0.5 0.5 0.5 0.1 0 102 61.5 3.9 2.3 0.2 0.1 0 0 0.1 0.1 0 0 0.1 0.3 0.2 0.2 1 1 0.2 0 1 0 0 0 0.3 0.3 0..1 0 1 1 0 0 0.5 1.1 1.3 1 0.7 2.8 0.3 0.5 0 0.1 0.1 0 7.3 8.9 5.3 4.8 9 1 6 2.6 4 . 8 9.7 10.4 1.9 0.3 197 Rhagium inquisitor (L inneaus) Rhizophagus pseudobrunneus Bousquet Rhizophagus remotus LeConte Rhyncolus brunneus Mannerhe im Rhyncolus macrops Buchanan Rhyncolus macrops Buchanan Rhyzophagus dimidiatus Mannerhe im Sacium lugubre LeConte Scierus annectans LeConte Scierus pubescens Swa ine Scolytus piceae (Swaine) Scolytus sp (unispinosus) LeConte Scolytus tsugae (Swaine) Scolytus tsugae (Swaine) Scolytus unispinosus LeConte Scotochroa basalis LeConte Sericoda quadripunctata (DeGeer ) Serralopalpus substriatus Ha ldeman Serralopalpus substriatus Ha ldeman Siagonium stacesmithi Hatch Siagonium stacesmithi Hatch sp#1 sp#11 sp#12 sp#13 sp#15 sp#3 sp#6 sp # 608 Sphaeriestes alternatus (LeConte) Sphaerites politus Duf tschmid Stenus bilineatus J . Sahlberg Stephanopachys substriatus (Paykul l ) Strictoleptura canadensis cribripennis (LeConte) Syneta albida LeConte Syneta pilosa W . J . Brown Syntomus americanus (Dejean) Tachinus elongatus Gyl lenhal Tachinus elongatus Gyl lenhal Teretruis montanus Horn Tetropium velutinum LeConte Thalycra mixta H. Howden Thanasimus undatulus (Say) Thanatophilus lapponicus (Herbst) Tomoxia borealis (LeConte) Trachypachus holmbergi Mannerhe im Trachysida a. aspera (LeConte) Triadhron lecontei Horn Tribolium audax Halstead Trichocellus cognatus (Gyl lenhal) Cerambyc idae 0 9 11 0 . 4 0 . 2 Rhizophagidae 0 0 0 0 . 4 0 Rhizophagidae 0.6 0.8 0.7 0.1 0.5 Curcul ionidae 0.2 0.2 0.5 0.2 0.3 Curcul ionidae 1.7 2.6 2.5 0.9 0.8 Curcul ionidae 1.6 3 3 0 . 2 1.2 Rhizophagidae 0 . 1 1 1 0 0 Cory lophidae 0 4.5 1.3 1.6 0 Scolyt idae 4 . 4 12 2 0 . 8 0 Scolyt idae 0.1 0.2 0 0 0 Scolyt idae 0 4.7 0.6 0.3 0.3 Scolyt idae 0 15.7 14.2 1.2 0.2 Scolyt idae 1.3 9.2 1 0.1 0 Scolyt idae 0 7 0 0 0 Scolyt idae 0.4 28 12.5 1.4 0.2 Melandry idae 0.1 0.4 0.7 0.5 0 Carabidae 0.1 1.2 0.5 0.6 0 Melandry idae 0.6 1.5 0.7 0.1 0 Melandry idae 0 . 6 2 0 0 0 Staphyl in idae 0.6 0.8 2.1 1 0.2 Staphyl in idae 0 3 3 0 . 4 0 . 2 Chrysomel idae 0 0.1 0.1 0 0 Nitidul idae 0 0.1 0.1 0 0 Nitidul idae 0 0.3 0.1 0 0 Nitidul idae 0 0.5 0.1 0 0 Nitidul idae 0 0.2 0.2 0 0 Nitidul idae 0 0.1 0.1 0.1 0 Nitidul idae 0 0.2 0 0 0 Scarabae idae 0 0 0.1 0.1 0 Salpingidae 0 0.4 0.1 0.2 0 Spaer i t idae 0 0 0.1 0.1 0 Staphyl in idae 0 0.8 0.3 0.2 0.2 Bostr ichidae 0 0.2 0.1 0 0 Cerambyc idae 0 0 1 1 2 . 6 5 . 6 Chrysomel idae 0.3 1.3 1.3 0 0.2 Chrysomel idae 0 0.6 0.4 0 0 Carab idae 0.1 0.6 0.5 0.5 0.3 Staphyl in idae 0.1 0.5 0 0 0 Staphyl in idae 0 . 3 2 1 0 . 2 0 Hister idae 0 0 0 0 . 8 0 . 2 Cerambyc idae 0.4 1.5 0.6 0.1 0 Nitidul idae 0 0.4 0.5 0 0 Cler idae 2 4 4 5 0 . 6 0 . 4 ' Silphidae 0 0.4 0.2 0.2 0 Mordel l idae 0 0.1 0.3 0.2 0.2 Carab idae 0 2.2 1.3 0.1 0.2 Cerambyc idae 0 . 3 1 3 0 0 . 4 Leiodidae 0 0.1 0.1 0 0 Tenebr ion idae 0.3 4.3 1.5 1.7 1 Carabidae 0 1.8 1.9 2.1 1.3 198 Trichocellus cognatus (Gyl lenhal) Trichodes ornatus hartwegianus A. Whi te Trichophya pilicornis (Gyl lenhal) Trypodendron Linneatum (Olivier) Trypodendron Linneatum (Olivier) Typhaea stercorea (L inneaus) Xestoleptura tibialis (LeConte) Xyleta laevigata (Hel lenius) Xyleta laevigata (Hel lenius) Xyletinus rotundicollis R.E. Whi te Xylotrechus longitarsis (Casey) /undulatus (Say) Carab idae 0 0 2 0 . 6 0 . 6 Cler idae 0 0 0.1 0.2 0 Staphyl in idae 0 1.2 0 .6 0 .2 0 Scolyt idae 1 2 7 4 4 0 1 3 4 8 5 3.4 1.3 Scolyt idae 1 3 5 9 8 2 1 0 . 2 Mycetophag idae 0 0.2 0 0 0 Cerambyc idae 0 0.2 0 .8 0.2 0 .2 Melandry idae 2 . 9 8 7 . 7 2 7 . 1 8.4 6 Melandry idae 0 . 4 8 2 70 6 . 2 7 . 4 Anobi idae 0 0 0.2 0 .2 0 Cerambyc idae 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 A b u n d a n c e / Site Species - Decreasing Then Increasing sharves w w CO (/) •9 y) Abundance Family i CL o C L o C L o C L o C L Acidota• crenata (Fabric ius) Staphyl in idae 0 . 3 0 0 0. 1.2 Bolitopunctus muricatulus Staphyl in idae 3 1 0 2. 2 Calopus angustus LeConte Oedemer inae 0 . 7 2 0 1.4 Corcodera (prob) longicornis (Kirby) Cerambyc idae 0 . 1 0 0 0 . 2 Codicaria n sp Lathridi idae 2 . 6 1 1 1.8 Cryptophagus sp# 1 Cryptophag idae 0.4 0.4 0.1 0.3 Cryptophagus sp#3 Cryptophag idae 0 . 7 0 0 0. 0 . 8 Ctenicera angusticollis (Mannerhe im) Elateridae 0 . 9 0 0 0. 1.2 Ctenicera mendax (LeConte) Elateridae 0 . 4 0 0 0 . 2 Ctenicera semimetallica (Walker) Elater idae 0 . 1 0 0 0. 1 Epuraea flavomaculata Makl in Nit idulidae 0.1 0.1 0 0.7 Eusphalerum spp (mostly pothos (Mannerhe im)) Staphyl in idae 6 . 9 6 1 3. 2.2 Evodinus monticola vancouveri Casey Cerambyc idae 2.7 0.2 0 0. 0.2 Hydnobius sp#3 Leiodidae 0 . 1 0 0 0. 0 . 2 Lordithon cascadensis (Malkin) Staphyl in idae 0 . 1 0 0 0. 0 . 6 Mycetoporus americanus Er ichson Staphyl in idae 0 . 1 0 0 0. 0 . 2 Mycetoporus rufohumoralis Staphyl in idae 1.9 1 0 0. 0 . 6 Odontosphindus clavicornis Casey Sphindidae 0 . 1 0 0 0. 0 . 8 Philonthinii spp Staphyl in idae 0 . 3 0 0 0. 0 . 4 Rhinosimus viridiaeneus Randal l Salp ingidae 2.1 1.8 0.7 1 Tachinus basalis Er ichson Staphyl in idae 0 . 7 0 0 0. 0 . 4 Tachinus frigidus Er ichson Staphyl in idae 0 . 1 0 0 0 . 2 Tetratoma concolor LeConte Tet ra tomidae 1.4 0 0 0 . 2 Trichochrous albedensis Blaisdell Melyr idae 0.4 0.2 0.5 0. 0.7 Trichochrous albedensis Blaisdell Melyr idae 2.9 0 0 0 . 4 Triplax dissimulator (Crotch) Erotyl idae 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 A b u n d a n c e / Site co CD £ CD .C CM CO Species - No Abundance Trend Family CD Q_ to o D. to o 0. to o D. to O a. Acmaeops p. protects (Kirby) Cerambyc idae 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 Mannerhe im Staphyl in idae 0.1 0 0.2 0.2 0 Aleocharinae (misc.spp) Staphyl in idae 0 1 0 0 0.2 Amara erratica (Dutschmid) Carab idae 0 1.2 0.5 1 0.2 Amara idahoana (Casey) Carab idae 0.1 0 0.1 0.1 0.2 Ampedus behrensi + phelpsi (Horn) Elater idae 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 Scrapt i idae 1.4 3 6 1 4 3 . 6 Anaspis sp#2 Scrapt i idae 0 1 3 0 0 . 4 Anisotoma globososa Hatch Leiodidae 0 . 4 0 1 1.2 0 . 4 Anotylus tetracarinatus (Block) Staphyl in idae 0 0 0.3 0 0.2 Anthaxia inornata (Randal l ) Buprest idae 0 . 1 0 1 0.2 0 Antherophagus sp# 1 Cryptophag idae 0.4 0.2 0.4 0.1 0.2 Aphodius fimetarius (L inneaus) Scarabae idae 0.8 0.1 0 0.1 0 Aphodius leopardus Horn Scarabaeidae 0 0 0.3 0 0.2 Atheta dentata Staphyl in idae 0 . 1 3 0 0 0 . 2 Athous nigropilis Motschulsky Elater idae 0.4 0 0.1 0 0 Atomairia sp# 13 Cryptophag idae 0 0 0.2 0 0.2 Atomaria sp#3 Cryptophag idae 0 0.2 0 0.1 0 Atrecus quadripennis (Casey) Staphyl in idae 0.1 0 0 0.1 0 Bius e s t r i a t u s (LeConte) Tenebr ion idae 0.3 0 0 0.2 0 Bradycellus lecontei Csiki Carabidae 0 0.3 0.2 0.8 0.3 Buprestis lyrata Casey Buprest idae 0 6.5 4.5 4.5 6.2 Buprestis nuttalli Kirby Buprest idae 0 0 0 0 0.6 Byturus unicolor Say Bytur idae 0 1 0 0 . 2 0 Caenocelis sp # 1 Cryptophag idae 0 . 3 0 2 0 . 6 1 Calathus advena (Leconte) Carab idae 0.1 0 0.3 0 0.2 Cephaloon tenuicorne LeConte Cephalo idae 0 . 6 0 2 0.2 1 Cis sp (fuscipes) Mell ie Ci idae 0.3 0.3 0 0.3 0 Clavilispinus rufescens (Hatch) Staphyl in idae 0 0.1 0 0.1 0 Coccinella septumpunctata L inneaus Coccinel l idae 0 0 4 5.8 3.2 Coccinella trifasciata perplexa Mulsant Coccinel l idae 0 1 1 0.8 2 Colon magnicolle Mannerhe im Leiodidae 0 0 1 0 0 . 4 Corcodera (prob) longicornis (Kirby) Cerambyc idae 0 0.2 0.1 0.3 0.2 Codiceus praetermissus (Fall) Tenebr ion idae 0 1 0 0 0.2 Codiceus tenuis (LeConte) Tenebr ion idae 0 0.1 0 0.1 0 201 Cortodera m. militaris (LeConte) Cossonus pacificus V a n Dyke Cryptophagus sp#2 Cryptophagus sp#4 Ctenicera bombycina (Germar ) Ctenicera comes (W . J . Brown) Ctenicera crestonensis (W . J . Brown) Ctenicera kendalli Kirby Ctenicera lutescens (Fa\\)/sagitticollis (Eschschol tz) Ctenicera nebraskensis (Bland) Ctenicera nigricollis (Bland) Ctenicera nitidula (LeConte) Ctenicera pudica(\NJ. Brown)+propola columbiana (Leconte) Ctenicera volitans (Eschschol tz) Curimopsis sp Cyphon concinnus (LeConte) Cydusa sp (subtestacea) (Gyl lenhal) Cytilus sp (alternatus) (Say) Cytilus sp (alternatus) (Say) Dendroctonus rufipennis (Kirby) Dendrophagus cygnaei Mannerhe im Dermestes talpinus Mannerhe im Dicerca tenebrosa (Kirby) Dolichocis manitoba Dury Dorcatoma (prob) americana # 87 Dyctyopterus spp Eanus decoratus (Mannerhe im) Eanus sp # 1 Enicmus tenuicornis LeConte Epiphanis cornutus Eschschol tz Epuraea (nr) populi Dodge Epuraea depressa Ernobius gentilis Fall Ernobius gentilis Fall Ernobius nigrans Fall Eupraea truncatella Mannerhe im Glischrochilus moratus W . J . Brown Grammoptera subargentata (Kirby) Grammoptera subargentata (Kirby) Harpalus fuscipalps Harpalus nigritarsis C.R. Sahlberg Hemicoelus carinatus (Say) Hydnobius sp#3 Hydroporus sp # 2 Hylastes longicollis Swa ine Hylastes longicollis Swa ine Hylastes ruber Swa ine Hylurgops reticulatus W o o d Hylurgops rugipennis Mannerhe im/F i tch Cerambyc idae 0 Curcul ionidae 0 Cryptophagidae 0.4 Cryptophagidae 0.7 Elateridae 0 . 1 Elateridae 0 . 1 Elateridae 0 Elater idae 0 Elateridae 0 . 3 Elateridae 1.4 Elateridae 1.7 Elateridae 0 Elateridae 5 . 1 Elateridae 2.6 Byrrhidae 0.1 Scirt idae 0 Leiodidae 0 Byrrhidae 0 Byrrhidae 0 Scolyt idae 0 . 6 Cucuj idae 0 . 3 Dermest idae 0 . 1 Buprest idae 0 Ciidae 0 Anobi idae 0.4 Lycidae 2.3 Elateridae 0 Elateridae 0 . 6 Lathridi idae 2 . 7 Eucnemidae 0.2 Nitidul idae 0 Nitidulidae 1.1 Anobi idae 0 Anobi idae 0 Anobi idae 0 Nitidulidae 0 Nitidulidae 0 Cerambyc idae 0 Cerambyc idae 0 . 1 Carabidae 0 Carabidae 0 Anobi idae 0 . 1 Leiodidae 0 Dytiscidae 0 Scolyt idae 0.3 Scolyt idae 0 . 1 Scolyt idae 7 . 3 Scolyt idae 0 Scolyt idae 0 4 0 0 0 . 6 2 0 0 . 2 0 0.2 0.2 0.2 0.2 0.6 0.8 0.5 0.2 0 0 1 0.2 0 1 0 0 0.5 0 0.3 0.3 0.2 0.4 0.1 0.7 1 0 0 . 2 0 1 0 5 . 2 2 4 . 6 8 0 1.8 1 0 . 2 2 0 0 . 6 0 . 2 4 3 1 2 0 . 2 2 8 . 6 1.9 1 2.3 1.3 0 0.1 0 0.2 0 0.2 0 0.2 0 0.1 0 0.3 0.3 0.5 0.2 0.7 1 0 0 0 . 2 0 1 0.4 0 0 2 0 . 2 1 0 0 0 . 2 0 5 0 1.6 0 . 4 0 1 0 0 . 2 0 0.1 0 0 0.7 2.4 1.2 0.7 1 0 0.2 0 1 0 0.2 0 0 3 1.8 1 0 0.1 0 0 0.2 0.1 0 0.2 0.1 0 0.2 0.2 0.4 0 0 0.2 1 0 0 0.6 0.1 0 0.2 0 2 0 0 . 2 0 0.2 0.1 0 0.3 0.4 0.1 0.7 0.2 0 1 0.4 1.2 0 0.2 0 0.2 0.1 0 0.1 0 0 0 0.2 0 0.1 0 0.2 0 0 0 0.2 0 0.8 0.4 0.8 0.3 1 0 0 0.4 2 6 1 3 . 2 7 0.2 0 0.1 0 0.8 0 0.1 0 202 Hypnoidus bicolor (Eschschol tz ) Ips latidens (LeConte) Ips pini (Say) Ips tridens (Mannerhe im) Ischnosoma sp (fibratum) Lacon rorulentus (LeConte) Lacon rorulentus (LeConte) Lasconotus complex LeConte Lasconotus intricatus Kraus Leiodes strigata (LeConte) Limonius aeger LeConte Limonius aeger LeConte Lordithon bimaculatus Lordithon fungicola Campbel l Lordithon poecilus (Mannerhe im) Lordithon t. thoracicus (Fabric ius) Megadhrus angulicollis Megatoma verigatta (Horn) Melanophila drummondi (Kirby) Micropeplus laticollis Makl in Micropeplus laticollis Makl in Molamba obesa Casey Monochamus spp Mulsantina picta (Randal l ) Mycetochara fraterna (Say) Mycetophagus distinctus Hatch Mycetoporus americanus Er ichson Myrmedophila americana (LeConte) Neanthophlax mirificus (Bland) Nudobius cephalus (Say) Olophrum consimile (Gyl lenhal) Omalium sp (foraminosum MaklinJ Orchesia (nr) castanea Orphilis subnitidus LeConte Orsodacne atra (Ahrens) Orsodacne sp Odhocis punctatus Casey Ostoma ferrugina (Linneaus) Oxytelus fuscipennis Mannerhe im Oxytelus fuscipennis Mannerhe im Paromalus mancus Casey Phausis rhombica Fender Philodrepa (?) Dropephylla sp (nrlongula) Philonthus politus (L inneaus) Phymatodes dimidiatus (Kirby) Phymatodes dimidiatus (Kirby) Pissodes fasciatus LeConte Pityogenes knetchteli Swa ine Pityogenes plagiatus (LeConte) Pityophthorus pseudotsugae Swa ine Elateridae 0 Scolyt idae 0 Scolyt idae 0 Scolyt idae 0 Staphyl in idae 0 Elateridae 0 Elater idae 0 Colydi idae 0 Colydi idae 0.1 Leiodidae 0 Elater idae 0 Elater idae 0 Staphyl in idae 0.3 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0 . 1 Dermest idae 0 . 7 Buprest idae 0 Staphyl in idae 0.1 Staphyl in idae 0 . 1 Cory lophidae 0 . 1 Cerambyc idae 0 Coccinel l idae 0.2 Tenebr ion idae 0.1 Mycetophag idae 0 . 4 Staphyl in idae 0.1 Cryptophag idae 0 Cerambyc idae 0.3 Staphyl in idae 0 Staphyl in idae 0 Staphyl in idae 0.2 Scrapt i idae 0 . 1 Dermest idae 0 Chrysomel idae 0 . 6 Chrysomel idae 0 . 1 Ciidae 0 . 3 Trogossi t idae 0.6 Staphyl in idae 0.2 Staphyl in idae 0 . 3 Hister idae 0 Lampyr idae 0 . 1 Staphyl in idae 0 . 4 Staphyl in idae 0.1 Cerambyc idae 0.5 Cerambyc idae 0 . 1 Curcul ionidae 0 . 7 Scolyt idae 0 Scolyt idae 0 Scolyt idae 0 0.5 0.7 0.2 0.5 0.2 0 0.1 0 5 1 0 0.2 2 1 0 0.2 0.1 0 0.1 0 2.4 0.9 0.4 2 1 2 0 0.2 0.2 0 0 0.2 0.1 0 0.1 0 1 0 0 . 2 0.2 0.3 0.3 0.2 0.8 1 0 0 . 2 0 0 0 0.1 0 0 1 0 . 2 0 . 4 0.1 0.2 0 0.2 1 0 0 0.2 1 0 0 . 4 0 1 3 0 0 . 2 0.2 1 0 0 . 4 0 0 0.3 0.1 0.2 0 0 0 . 2 0 1 0 0 0.6 4 0 0 . 2 0 0.3 0.2 0.1 0.2 0 0.3 0.1 0 0 0 1 0 . 4 0 0.3 0.3 0 0 1 0.2 0.8 2.8 1.4 1.4 5.7 0 1 0.2 0 . 6 0 0 0.2 0 0.1 0.1 0.2 0 0 0 0 . 4 0 0.4 0.2 0.1 0.2 0 0 4 . 2 . 0 . 4 0 0 0 . 4 0 0 2 0 . 2 0 0.1 0.4 0.6 0.2 0 0.5 1.6 0.8 0 1 0 . 6 1.2 4 6 0 0 . 2 0 0 0 . 2 0 2 4 1.6 4 0 0.1 0 0 0.6 0.5 0.2 0.5 0 1 0 0 3 0 0 . 4 0.2 0.7 0 0.1 0 2 0 0 . 2 0.2 2 0 0 . 2 0 203 Pityopthorus aquilus B lackman (+ aplanatus) Platydema spit 1 Plegaderus sayi Marseul Podabrus piniphilus (Eschschol tz) Podabrus scaber LeConte Podabrus scaber LeConte Poliaenus oregonus (LeConte) Polygraphus convexifrons W o o d Pseudohylesinus nebulosus LeConte Pseudopsis sp Ptinus californicus Pic Pygoleptura n. nigrella (Say) Quediini spp Quedius plagiatus Mannerhe im Quedius s. spelaeus Horn Rhinosimus viridiaeneus Randal l Rhizophagus remotus LeConte Sacium lugubre LeConte Salebius nr minax Scaphisoma castaneum Motschulsky Scolytus sp (unispinosus) LeConte Scolytus unispinosus LeConte Scotochroa basalis LeConte Scymnus sp Semanotus ligneus (Casey) Semijulistus ater (LeConte) Silis d. difficilis LeConte Sitona cylindricollis (Fahraeus) sp#2 sp#3 sp#4 s p # 9 Spondylis upiformis Mannerhe im Staphylinus pleuralis LeConte Staphylinus pleuralis LeConte Stephostethus liratus (LeConte) Syneta albida LeConte Syneta hamata Horn Syntomium grahami Hatch Tachinus basalis Er ichson Tachinus thruppi Hatch Tachyporus sp (canadensis Campbel l ) Thalycra mixta H. Howden Trachypachus holmbergi Mannerhe im Tragosoma depsarium (L inneaus) Triadhron lecontei Horn Trichophya pilicornis (Gyl lenhal) Triplax antica LeConte Triplax antica LeConte Triplax californica LeConte Scolyt idae 0 Tenebr ion idae 0 Histeridae 0 Canthar idae 0 . 6 Canthar idae 0 Canthar idae 0 Cerambyc idae 0 Scolyt idae 0 . 1 Scolyt idae 4 . 7 Staphyl in idae 0.1 Ptinidae 0 Cerambyc idae 0.1 Staphyl in idae 0 . 1 Staphyl in idae 1.6 Staphyl in idae 0.1 Salpingidae 0 . 1 Rhizophagidae 0 . 6 Cory lophidae 0 . 1 Cryptophagidae 0.4 Scaphidi idae 0.1 Scolyt idae 0 . 1 Scolyt idae 0 . 1 Melandry idae 0 . 6 Coccinel l idae 0 Cerambyc idae 0 Melyr idae 0 Canthar idae 0.1 Curcul ionidae 0 Ciidae 0 Ciidae 0 Nitidulidae 0 Nitidulidae 0 Cerambyc idae 3 . 3 Staphyl in idae 0.1 Staphyl in idae 0 . 1 Lathridi idae 0 . 4 Chrysomel idae 0 . 6 Chrysomel idae 0 Staphyl in idae 0 Staphyl in idae 0.7 Staphyl in idae 0.1 Staphyl in idae 0 Nitidulidae 0 . 4 Carab idae 0 Cerambyc idae 0 Leiodidae 0 . 1 Staphyl in idae 0 . 1 Erotyl idae 0 Erotyl idae 0 . 1 Erotyl idae 0.4 0.2 0.1 0.1 0.2 0 0 0.2 0 0.1 0.1 0 0.2 3 0 0 . 4 0 . 6 0 0.2 0 0.2 2 0 0 0.2 0.2 0.1 0 0.2 0 0 0 . 2 0 6 4 7 0 1.6 1.2 0 0.1 0.1 0 0.2 0.1 0.2 0.2 0.6 0.5 0.9 0.7 0 0 0 . 4 0 3 0 0 . 2 0 0 0.1 0 0 0 0 0 0 . 4 0 0 0 . 4 0 . 2 1 0 0 . 4 0 . 4 0 0.2 0 0 0.4 0.3 0.9 0.7 6 0 1.4 0 . 8 5 0 1.6 1.6 1 0 0 . 6 0 . 2 0 0.1 0 0.3 1 0 0 0 . 2 3.3 3.2 3.6 2.3 0.1 0 0.1 0 0 0.1 0 0.2 0.1 0 0.1 0 0.3 0 0.1 0.2 0.2 0 0 0.2 0.4 0.1 0 0.3 4 0 1 0 . 6 0.5 0.7 0.5 0.7 1 0 0 . 4 0.2 0 0 1 0 0 1 0 . 6 0 0.1 0 0 0.7 0 0.2 0 0.2 0.1 0.1 0.1 0.3 0 0.1 0.2 0 0 0.1 0 0.3 0 0 0 . 2 0 1 0 0 . 2 0 0.1 0.3 0.1 0.3 1 0 0 0.2 1 0 0 . 6 0 0.1 0 0 0.2 0 0 0 . 2 0 1 0.9 1.8 1.3 204 Triplax californica LeConte Trixagus sp Trypodendron betulae Swa ine Trypophloeus populi Hopkins Tychius picirostris (Fabr icus) Upis ceramboides (L inneaus) Utobium elegans (Horn) Xestoleptura tibialis (LeConte) Xylotrechus longitarsis (Casey) /undulatus (Say) Zilora occidentalis Mank Zyras sp. Erotyl idae 0 . 7 1 0 0 . 8 1.2 Throsc idae 0 0.3 0 0.3 0.3 Scolyt idae 0 0 0 0 .2 0 Scolyt idae 0 0.2 0 1.4 0 Curcul ionidae 0 0 0.1 0 0.2 Tenebr ion idae 0 0.2 0.1 0.2 0.2 Anobi idae 0.1 0 0.1 0.1 0 Cerambyc idae 0 1 0 0 . 4 0 Cerambyc idae 0 . 1 0 1 1.2 0 . 4 Melandry idae 0.2 0 0.1 0 0 .2 Staphyl in idae 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 preharvest Post 1 Post 2 Post 3 Post 4/5 sp # 296 Nitidulidae 0.1 0 0 0 0 sp # 299 Nitidulidae 0 0 0.1 0 0 Adalia bipunctata (L inneaus) Coccinel l idae 0 0 0 0 0.2 Aegialia rufescens Horn Scarabae idae 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 Gravenhors t Staphyl in idae 0 0.1 0 0 0 Aleochara suffosa (Casey) Staphyl in idae 0 0 0 0.2 0 Aleochara villosa Mannerhe im Staphyl in idae 0 0 0 0 0.2 Amara apricaria (Paykul l ) Carab idae 0 0.1 0 0 0 Amara apricaria (Paykul l ) Carab idae 0 0 0 0 0.2 Amara sinuosa (Casey) Carab idae 0 0.1 0 0 0 Anthobium reflexicolle Casey Staphyl in idae 0 0 0 0.1 0 Anthrenus pimpinellae Fabr icus Dermest idae 0 0 0.1 0 0 Aphodius distinctus (O.F. Mul ler) Scarabaeidae 0 0 0.1 0 0 Aphodius haemorrhoidalis/pectoralis (L inneaus) /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) Staphyl in idae 0 1 0 0 0 Attalus sp Melyr idae 0 0.1 0 0 0 Bembidion canadianum Casey Carab idae 0 0 0 0 0.2 Bembidion quadrimaculatum (LeConte) Carab idae 0 0 0 0 0.2 Bembidion timidum (LeConte) Carab idae 0 0.1 0 0 0 Bhyrrhus sp Byrrhidae 0.1 0 0 0 0 Bledius ruficornis LeConte Staphyl in idae 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 (L inneaus) Chrysomel idae 0 0 0 0.1 0 Bryophacis canadensis Staphyl in idae 0.1 0 0 0 0 Bryophacis spp Staphyl in idae 0 0 0 0 0.2 Calathus advena (Leconte) Carab idae 0.1 0 0 0 0 Canifa sp# 1 Melandry idae 0 0 0 0.2 0 Cardiophorus (prob) tenebrosus LeConte Elater idae 0 0 0 0.1 0 Carphoborus vandykei Bruck Scolyt idae 0 0 0 0.2 0 Cercyon sp#3 (tolfino) Hatch Hydrophi l idae 0 0 0 0 0.2 206 Cerycon sp#1 (herceus frigidus) Smetana Hydrophi l idae 0 0 0 0.2 0 Ceutorhynchus erysimi (Fabr ic ius) Curcul ionidae 0 0 0 0 0.2 Ceutorhynchus punctiger Gyl lenhal Curcul ionidae 0 0 0 0 0.2 Cimberis (prob) turbans Nemonych idae 0 0 0.1 0 0 Cis angustus Hatch Ci idae 0 0 0 0 0.2 Cis sp (fuscipes) Mell ie Ci idae 0 0 0 0.2 0 Clytus sp Cerambyc idae 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 Lathridi idae 0 0.1 0 0 0 Corticarina (prob) cavicollis (Mannerhe im) Lathridi idae 0 0 0.1 0 0 Creophilus maxillosus (L inneaus) Staphyl in idae 0 0 0.1 0 0 Crepidodera sp Chrysomel idae 0 0 0 0 0.2 Cryphalus ruficollis Hopkins Scolyt idae 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 (Mannerhe im) Pyrochro idae 0 0 0.1 0 0 Dermestes lardarius L inneaus Dermest idae 0 0 0 0 0.2 Dermestes sp Dermest idae 0 0 0 0.1 0 Desmatogaster subconnata (Fall) Anobi idae 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 Scarabae idae 0 0 0 0.1 0 Diplotaxis brevicollis LeConte Scarabae idae 0 0 0 0 0.2 Dorcatoma (prob) americana # 87 Anobi idae 0.1 0 0 0 0 Dryocetes betulae Hopkins Scolyt idae 0.1 0 0 0 0 Dryocetes caryi/schelti Hopk ins /Swaine Scolyt idae 0 0 1 0 0 Eanus decoratus (Mannerhe im) Elateridae 0 0 0.1 0 0 Elaphrus clairvillei Kirby Carab idae 0 0 0 0.1 0 Elaphrus clairvillei Kirby Carab idae 0 0 0 0 0.2 Eleates explanatus Casey Tenebr ion idae 0.1 0 0 0 0 Enoclerus sphegeus (Fabr icus) Cler idae 0 0 0 0.2 0 Epiphanis cornutus Eschschol tz Eucnemidae 0 0 0 0 0.2 Eucinetus (nr.) oviformis LeConte Eucinet idae 0 0.1 0 0 0 Eucnecosum tenue (LeConte) Staphyl in idae 0 0 0 0 0.2 Eupraea terminalis Mannerhe im Nit idulidae 0.1 0 0 0 0 Glischrochilus confluentus (Say) Nit idul idae 0 0 0 0 0.2 Glischrochilus moratus W . J . Brown Nit idul idae 0 0 0 0 0.2 Gymnusa atra Casey Staphyl in idae 0 0 0.1 0 0 Gymnusa pseudovariegata K l imaszewski Staphyl in idae 0 0 0.1 0 0 Gymnusa pseudovariegata K l imaszewski Staphyl in idae 0 0 0 0.2 0 Gymnusa sp (grandiceps Casey) Staphyl in idae 0 0 0.1 0 0 Gyrohypnus fracticornis (O.F. Muller) Staphyl in idae 0 0 0.1 0 0 207 Hallomenus sp Scrapt i idae 0 0.1 0 0 0 Harpalus obnixus Casey Carab idae 0 0 0.1 0 0 Harpalus opacipennis (Ha ldeman) Carab idae 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 S m e t a n a Staphyl in idae 0 0.1 0 0 0 Hippodamia tredecimpunctata (Say) Coccinel l idae 0 0.1 0 0 0 Hippuriphila sp Chrysomel idae 0 0 0 0 0.2 Hydaticus aruspex Clark Dyt iscidae 0 0 0 0 0.2 Hygrotus impressopunctatus (Schal ler) Dyt isc idae 0 0 0 0.1 0 Hypnoides impressicollis (Mannerhe im) Elateridae 0 0.1 0 0 0 Ips latidens (LeConte) Scolyt idae 0.1 0 0 0 0 Ips mexicanus (Hopkins) Scolyt idae 0 0.1 0 0 0 Isomera (nr) comstoki Papp Allecul idae 0 0 0.1 0 0 Laccobius sp Hydrophi l idae 0 0 0 0.1 0 Laemophloeus biguttatus (Say) Cucuj idae 0 0 0 0 0.2 Laricobius laticollis Fall Derodont idae 0 0.1 0 0 0 Lathridius ventralis Lathridi idae 0 0 0 0.2 0 Lathrobium negrum LeConte Staphyl in idae 0.1 0 0 0 0 Lebia moesta LeConte Carab idae 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) Coccinel l idae 0 0 0.1 0 0 Magdalis alutacea LeConte Curcul ionidae 0.1 0 0 0 0 Malthodes sp. Canthar idae 0 0 0 0 0.2 Malthodes sp. Canthar idae 0 0 0 0.2 0 Margarinotus rectus (Casey) Hister idae 0 0.1 0 0 0 Micetoporus sp Staphyl in idae 0.1 0 0 0 0 Micetoporus sp Staphyl in idae 0 0 0 0 0.2 Mycetochara fraterna (Say) Tenebr ion idae 0 0 0 0.2 0 Mycetophagus tenuifasciatus Horn Mycetophag idae 0 0 0.1 0 0 Mycetophagus tenuifasciatus Horn Mycetophag idae 0 0 0 0 0.2 Mycetoporus brunneus (Marsham) Staphyl in idae 0 0 0 0 0.2 Mycetoporus rugosus Hatch Staphyl in idae 0 0 0 0.2 0 Myremcocephalus arizonicus (Casey) Staphyl in idae 0 0.1 0 0 0 Neohypdonas tumescens (LeConte) Elater idae 0 0 0 0.1 0 Neohypnus obscurus (Er ichson) Staphyl in idae 0 0.1 0 0 0 Neohypnus obscurus (Er ichson) Staphyl in idae 0 0 0 0 0.2 Notiophilus aquaticus (L inneaus) 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) Staphyl in idae 0 0.1 0 0 0 Octotemnus denudatus Casey Ci idae 0 0 0.1 0 0 Octotemnus denudatus Casey Ci idae 0 0 0 0.2 0 Oiceoptoma noveboracense (Forster) Si lphidae 0 0 0 0.1 0 Omalium n. sp. Staphyl in idae 0 0 0.1 0 0 Omalium sp# 1 Staphyl in idae 0 1 0 0 0 Omalium spp. Staphyl in idae 0.1 0 0 0 0 Orchesia ornata Scrapt i idae 0 0 0 0.1 0 Orsodacne sp Chrysomel idae 0 0 0 0 0.2 208 Orus sp Staphyl in idae 0 0 0 0.1 0 Pachybrachis melanostictus Suffrain Chrysomel idae 0 0 0 0.1 0 Pachybrachis melanostictus Suffrain Chrysomel idae 0 0 0 0.2 0 Phaedon laevigatus (Duf tschmid) Chrysomel idae 0 0 0 0.2 0 Philonthus concinnus (Gravenhors t ) Staphyl in idae 0 0.1 0 0 0 Philonthus couleensis Hatch Staphyl in idae 0 0.1 0 0 0 Philonthus crotchi Horn Staphyl in idae 0 0.1 0 0 0 Philonthus varians (Paykul l ) Staphyl in idae 0 0.1 0 0 0 Phloeopra sp. Staphyl in idae 0 0.1 0 0 0 Phloeosinus pini Swa ine Scolyt idae 0 0.1 0 0 0 Phloeotribus lecontei Schedl /picea Swa ine Scolyt idae 0.1 0 0 0 0 Phymatodes (nr) fulgidus Hopping Cerambyc idae 0 0 1 0 0 Phymatodes maculicollis LeConte Cerambyc idae 0.1 0 0 0 0 Pidonia scripta (LeConte) Cerambyc idae 0.1 0 0 0 0 Pissodes (nr) fiskei Hopkins Curcul ionidae 0 1 0 0 0 Pissodes striatulus dubius Randal l Curcul ionidae 0 1 0 0 0 Pityomacer pix Kuschel Nemonych idae 0 0.1 0 0 0 Pityopthorus sp Scolyt idae 0 0.1 0 0 0 Plateumaris rufa (Say) Chrysomel idae 0 0 0 0.2 0 Platydema americanum Caste lnau & Brulle Tenebr ion idae 0 0.1 0 0 0 Platydema sp # 1 Tenebr ion idae 0 1 0 0 0 Platysoma leconti Marseul Hister idae 0 0 0 0.2 0 Plegaderus setulosus Ross Hister idae 0 0.1 0 0 0 Podabrus fissilis Fall Canthar idae 0 0 0 0.1 0 Podabrus sp # 613 Canthar idae 0 0 0.1 0 0 Pogonocherus mixtus Ha ldeman Cerambyc idae 0 0.1 0 0 0 Proteinus sp. Staphyl in idae 0 0.1 0 0 0 Pselaphus bellax Casey Pselaphidae 0 0.1 0 0 0 Pseudohadrotoma sp (perversa) (Fall) Dermest idae 0 1 0 0 0 Psyllobora vigintimaculata (Say) Coccinel l idae 0.1 0 0 0 0 Pterostichus adstrictus Eschschol tz Carabidae 0.1 0 0 0 0 Ptilinus lobatus/basalis Casey /Leconte Anobi idae 0 0 0 0 0.2 Quedius erythrogaster Mannerhe im Staphyl in idae 0.1 0 0 0 0 Quedius pediculus (Nordmann) Staphyl in idae 0 0 0 0.1 0 Quedius transparens Motschulsky Staphyl in idae 0 0 1 0 0 Rhantus binotatus (Harr is) Dyt iscidae 0 0.1 0 0 0 Rhyncolus brunneus Mannerhe im Curcul ionidae 0 0 0 0 0.2 Saprinus lugens Er ichson Histeridae 0 0.1 0 0 0 Scaphidema aeneolum (LeConte) Tenebr ion idae 0 0 0.1 0 0 Scaphium sp Scaphidi idae 0 0 0 0.2 0 Scolytus opacus B lackman Scolyt idae 0 0.1 0 0 0 Scolytus subscaber LeConte Scolyt idae 0 0.1 0 0 0 Scraptiidae sp#3 Scrapt i idae 0 0 0 0 0.2 Semanotus ligneus (Casey) Cerambyc idae 0 0.1 0 0 0 Sepedophilus littoreus (L inneaus) Staphyl in idae 0 0 0 0.2 0 Sericoda quadripunctata (DeGeer ) Carab idae 0 0 0 0 0.2 Sericus brunneus Elateridae 0 1 0 0 0 Silis d. difficilis LeConte Canthar idae 0 0 0 0.2 0 Sitona lineellus (Bonsdorf f ) Curcul ionidae 0 0 0.1 0 0 Sitona lineellus (Bonsdorf f ) Curcul ionidae 0 0 0 0.2 0 209 Sonoma parviceps (Makl in) Staphyl in idae 0 0 0 0 0.2 sp#1 Chysomel idae 0 0 0.1 0 0 sp#1 Anobi idae 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 Nitidul idae 0 1 0 0 0 sp#3 Ciidae 0 0 0 0.2 0 sp#4 Nitidul idae 0 0 0 0.2 0 sp # 573 Throsc idae 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 Scarabae idae 0 0 0 0 0.2 sp # 621 Dyt iscidae 0 0 0 0.2 0 sp # 623 Dyt iscidae 0 0 0.1 0 0 sp # 624 Dyt iscidae 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 Phalacr idae 0 0 0.1 0 0 sp#1 Phalacr idae 0 0 0 0 0.2 Sphaeridium bipustulatum Fabric ius Hydrophi l idae 0 0 0.1 0 0 Sphaeriestes sp. Salpingidae 0 0 0.1 0 0 Stenolophus fuliginosus Dejean Carab idae 0 0.1 0 0 0 Stenotrachelus aeneus (Fabr ic ius) Stenotrachel idae 0 0 0 0 0.2 Stenus juno Paykul l Staphyl in idae 0 0 0 0.1 0 Stenus plicipennis (Casey) Staphyl in idae 0 0 0.1 0 0 Stephostethus cinnamopterus (Mannerhe im) Lathridi idae 0 0.1 0 0 0 Syneta hamata Horn Chrysomel idae 0.1 0 0 0 0 Synuchus impunctatus (Say) Carabidae 0 0 0.1 0 0 Tachinus nigricornis Mannerhe im Staphyl in idae 0.1 0 0 0 0 Tachinus nigricornis Mannerhe im Staphyl in idae 0.1 0 0 0 0 Tachinus vergatus Campbel l Staphyl in idae 0 0 0 0.2 0 Tachyporus sp Staphyl in idae 0 0 0 0.1 0 Tachyporus sp Staphyl in idae 0.1 0 0 0 0 Tachyporus sp (lecontei Campbel lJ Staphyl in idae 0 0 0 0.1 0 Tachyta angulata Casey Carabidae 0 0 0 0 0.2 Thanatophilus lapponicus (Herbst) Si lphidae 0 0 1 0 0 Tragosoma depsarium (L inneaus) Cerambyc idae 0 0 0 0 0.2 Triplax dissimulator (Crotch) Erotyl idae 0.1 0 0 0 0 Xestocis sp Ciidae 0 0 0 0 0.2 Zilora occidentalis Mank Melandry idae 0.1 0 0 0 0 Zyras sp. Staphyl in idae 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 I V - 1 . Diversity as measured by Species Richness (S) f rom baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps were baited with componen ts MCOL, seudenol , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. B a i t e d / P r e h a r v e s t P o s t 1 P o s t 2 Post 3 Post 4/5 Contro l Baited 65.00 + 15.46 1 24.80 + 23.25 1 45.40 + 25.67 98.80 + 16.40 97.6 + 6.34 Contro l 53.60 + 12.56 1 52.00 113.00 88.20 + 18.30 92.4 + 13.42 Table IV-2. Diversity as measured by M a r g a l e f s r ichness index (d) f rom baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone t raps baited with c o m p o n e n t s M C O L , seudeno l , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. B a i t e d / P r e h a r v e s t P o s t - | P o s t 2 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) f rom baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudeno l , and frontal in known to aggregate Douglas-f i r beetles. Post 1 and post 2 unbai ted data represent a single t rapping site. Baited / Contro l Preharvest Post 1 Post 2 Post 3 Post 4/5 Baited 0.13 + 0.05 0.13 + 0.04 0.12 + 0.03 0.18 + 0.05 0.38 + 0.16 Contro l 0.85 + 0.05 0.54 0.81 0.85 + 0.02 0.80 + 0.04 211 Table IV-4. Diversity as measured by Bril louin index f rom baited and unbai ted pheromone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudeno l , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted da ta represent a single t rapping site. Bai ted / Contro l Preharvest Post 1 Post 2 Post 3 Post 4/5 Bai ted 0.53 + 0.23 0.61 + 0.20 0.60 + 0.14 0 . 7 9 + 0.24 1 . 6 3 + 0.71 Contro l 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 ' i 0 ) f rom baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 , h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudenol , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. ^ a i t f d . / Preharvest Post 1 Post 2 Post 3 Post 4/5 Contro l Baited 0.23 + 0.10 0.28 + 0.09 0.27 + 0.06 0.36 + 0.11 0 . 7 5 + 0 . 3 4 Contro l 1.46 + 0.14 1.09 1.67 1.64 + 0.06 1.57 + 0.06 Tab le IV-6. Diversity as measured by S impson 's dominance index ( 1 - ) f r o m baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudenol , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. Baited / Contro l Preharvest Post 1 Post 2 Post 3 Post 4/5 Baited 0.24 + 0.16 0.17 + 0.07 0.19 + 0.07 0.23 + 0.07 0.51 + 0.22 Contro l 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 ( ) f rom baited and unbai ted phe romone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudeno l , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. Bai ted / Contro l Preharvest Post 1 Post 2 Post 3 Post 4/5 Bai ted 9.50 + 2.34 21.31 + 3.21 22.46 + 3.54 20.27 + 4.30 25.01 + 7.26 Contro l 29.28 + 7.52 36.04 46.60 44.67 + 6.56 41.03 + 6.26 Table IV-8. Diversity as measured by Taxomon ic Diversity index ( ) f rom baited and unbai ted phe romone trapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudenol , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. Bai ted / Control Preharvest Post 1 Post 2 Post 3 Post 4/5 Baited 14.22 + 7.68 11.29 + 4.09 11.85 + 3.28 1 6.47 + 5.38 36.04 + 15.66 Contro l 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 Dist inctness index ( *) f rom baited and unbai ted pheromone t rapping sites in Douglas-f i r habitat, preharvest through 4 / 5 t h season post harvest condi t ions ( = 0.05). Pheromone traps baited with componen ts M C O L , seudenol , and frontal in known to aggregate Douglas-f i r beet les. Post 1 and post 2 unbai ted data represent a single t rapping site. B a i t e d / p r e h a r v e s t P o s t 1 P o s t 2 Control Post 3 Post 4/5 Baited 63.01 + 8.65 65.73 + 1.98 64.33 + 6.89 71.51 + 0.67 71.49 + 0.96 Contro l 71.24 + 0.95 63.00 73.17 70.03 + 1.41 66.16 + 2.13 213 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
https://iiif.library.ubc.ca/presentation/dsp.831.1-0090382/manifest

Comment

Related Items