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Responses of ground beetle (Coleoptera: carabidae_ species and assemblages to forest practices in the… Jarrett, Jeffrey R. 2003

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RESPONSES OF GROUND B E E T L E (COLEOPTERA: CARABIDAE) S P E C I E S A N D A S S E M B L A G E S T O F O R E S T P R A C T I C E S IN T H E INTERIOR DOUGLAS-FIR FORESTS OF BRITISH COLUMBIA by  J E F F R E Y R. J A R R E T T B . Sc. University of V i c t o r i a , 1992  A THESIS S U B M I T T E D IN P A R T I A L F U L F I L L M E N T O F THE REQUIREMENTS FOR THE DEGREE OF MASTERS OF SCIENCE in  T H E F A C U L T Y OF G R A D U A T E STUDIES (DEPARTMENT OF ZOOLOGY)  W e accept this thesis as c o n f o r m i n g to the required s t a n d a r d  THE UNIVERSITY OF BRITISH COLUMBIA December, 2003 © Jeffrey R. J a r r e t t , 2003  Library Authorization  In presenting this thesis in partial fulfillment of the requirements for a n a d v a n c e d d e g r e e at the University of British C o l u m b i a , I a g r e e that the Library shall m a k e it freely available for reference a n d study! I further a g r e e that p e r m i s s i o n for e x t e n s i v e c o p y i n g of this thesis for scholarly p u r p o s e s m a y be granted by the h e a d of my department or by his or her representatives. It is u n d e r s t o o d that c o p y i n g or publication of this thesis for financial gain shall not be allowed without my written p e r m i s s i o n .  Date (dd/mm/yyyy)  N a m e of A u t h o r (please print)  Degree:  MOLSH^S  D e p a r t m e n t of  oi  _&:ie<Ac-e_  ?LD<3  T h e University of British C o l u m b i a Vancouver, B C  Canada  Year:  e_e>cO  ABSTRACT  The Opax Mountain Silvicultural Systems Project was initiated in 1993 to address concerns over the widespread use of uniform stand-level partial cutting in the dry Douglas-fir forests of BC's Southern Interior. Various alternative harvesting methods were tested, and responses of several wildlife indicator groups were measured. The following study examined responses of ground beetle (Coleoptera: Carabidae) species and assemblages using a variety of diversity indices. The alternative harvesting practices were: 1) 20% and 2) 50% removal using individual-tree selection (I.T.S.); 3) 20% and 4) 50% removal using patch-cuts of 0.1, 0.4, and 1.6 ha; 5) 35% removal using 50% I.T.S. on 70% of the treatment area, leaving 30% in reserves; and 6) uncut controls. Results showed that species richness, evenness, and heterogeneity were greater in logged treatments. However, only in the heterogeneity values of Mud Lake assemblages (1996/1997) was there a general increase in diversity that accompanied either an increase in percent forest removal, or a change from control to I.T.S. to patch-cut methods of harvest. Other than taxonomic distinctness, no other distinctness index decreased in logged treatments, and none of the distinctness indices displayed any trend in decreasing with an increasing percentage of forest removal, or to a change from control to I.T.S. to patch-cut methods of harvest. In multivariate analysis, carabid assemblages within each site (Mud Lake and Opax Mountain.) showed patch-cut treatments to cluster/map together and I.T.S. treatments to cluster/map together. Dominance structure tables generally showed that carabid assemblages from lower percent removal treatments, as well as from I.T.S. methods, to most resemble dominance structure found in control  ii  blocks. Species analysis showed that carabid species respond to logging in a variety of ways. Because every method of harvest evidently benefited some species at the expense of others, no one treatment appears sufficient for the whole carabid community. Instead, a mix of harvesting methods that maintained the greatest number of all native forest species, including sensitive species, and over an indefinite period of time, would be the best strategy for the preservation of carabid biodiversity.  in  T A B L E OF CONTENTS Abstract  ii  Table of contents  iv  List of Tables  vii  List of Figures  •  Acknowledgments  ix xiii  Introduction  1  Forest biodiversity  1  Definition/Importance  1  Biodiversity Numbers  2  Threats to Forest Biodiversity  2  Conservation of Biodiversity  3  Opax Mountain Silvicultural Systems Research Project.....  5  Carabidae  6  Diversity, distribution, biology and ecology  6  Endangered carabids  9  Indicators and Indices  12  Opax Carabids  13  Objectives and Hypothesis  16  Materials and Methods  18  Site location and description  18  Logging manipulations  21  iv  Sampling design  22  Arthropod sampling technique  23  Sorting and identification  24  Data analysis  25  Species richness  26  Evenness  26  Heterogeneity  27  Taxonomic distinctness  28  Multivariate analysis  29  Dominance structure  29  Species analysis  30  Statistical tests  30  Results  32  Species richness  51  Whittaker plots  54  Evenness  60  Alpha index  63  Shannon-Wiener index  67  Simpson's index  68  Taxonomic distinctness  73  Average taxonomic distinctness  77  Variation in taxonomic distinctness  80  Cluster analysis  87  V  Multidimensional scaling  88  Dominance structure  93  Species analysis  97  Discussion  107  Mud Lake and Opax Mountain  109  Spring vs. entire year sampling  112  Species richness  112  Evenness  114  Heterogeneity  116  Taxonomic distinctness  119  Multivariate analysis  122  Dominance structure  124  Species analysis  125  Conclusions  134  Literature Cited  137  Appendices  151  VI  LIST O F TABLES  Table 1: Total number of specimens caught in all experimental treatments at the Opax Mountain Silvicultural Systems Project site  38  Table 2: Total number of specimens caught in all experimental treatments at the Mud Lake site  40  Table 3: Total number of specimens caught in all experimental treatments at the Opax Mountain site  42  Table 4: Total number of specimens caught in all experimental treatments (spring samples) at the Opax Mountain Silvicultural Systems Project site  44  Table 5: Total number of specimens caught in all experimental treatments (spring samples) at the Mud Lake site  46  Table 6: Total number of specimens caught in all experimental treatments (spring samples) at the Opax Mountain site  48  Table 7: Abundance, species number, species richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Mud Lake, based on total trap numbers for spring samples, 1995-1997  50  Table 8: Abundance, species number, species richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Mud Lake, based on total trap numbers for the entire season, 1993-1997  50  Table 9: Abundance, species number, species richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Opax Mountain, based on total trap numbers for spring samples, 1995-1997  50  Table 10: Abundance, species number, species richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Opax Mountain, based on total trap numbers for the entire season, 1993-1997  50  Table 11: Dominance position (and percent abundance) of the major species in the experimental treatments at the Mud Lake site, based on spring samples, 1995-1997  95  Table 12: Dominance position (and percent abundance) of the major species in the experimental treatments at the Mud Lake site, based on complete year samples, 1993-1997  95  vii  Table 13: Dominance position (and percent abundance) of the major species in the experimental treatments at the Opax Mountain site, based on spring samples, 1995-1997 Table 14: Dominance position (and percent abundance) of the major species in the experimental treatments at the Opax Mountain site, based on complete year samples, 1993-1997  viii  LIST O F FIGURES  Figure 1: The Interior Douglas-fir biogeoclimatic zone (IDF) of British Columbia, and the location of the Opax Mountain Silvicultural Systems Project site  18  Figure 2: Layout of the Opax Mountain Silvicultural Systems Project site, showing the six experimental treatments at both the Mud Lake and Opax Mountain site  22  Figure 3: Design of pitfall trapping scheme at the Opax Mountain Silvicultural Systems Project site: (a) individual pitfall trap with cover; (b) trap circle of five traps; (c) two sets of five circles per block  23  Figure 4: Corrected species richness (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain) spring samples  53  Figure 5: Corrected species richness (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain) spring samples  53  Figure 6: Corrected species richness (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain) spring samples  54  Figure 7: Whittaker plots of rank/abundance for Mud Lake 1995 season, spring samples  57  Figure 8: Whittaker plots of rank/abundance for Opax Mountain 1995 season, spring samples  57  Figure 9: Whittaker plots of rank/abundance for Mud Lake 1996 season, spring samples  58  Figure 10: Whittaker plots of rank/abundance for Mud Lake 1996 season, spring samples  58  Figure 11: Whittaker plots of rank/abundance for Opax Mountain 1997 season, spring samples  59  Figure 12: Whittaker plots of rank/abundance for Mud Lake 1997 season, spring samples  59  Figure 13: Evenness (Pielou's J') values for 1995 season (Mud Lake and Opax Mountain) spring samples  61  Figure 14: Evenness (Pielou's J') values for 1996 season (Mud Lake and Opax Mountain) spring samples  61  IX  Figure 15: Evenness (Pielou's J') values for 1997 season (Mud Lake and Opax Mountain) spring samples  62  Figure 16: Alpha index values (+/- s.e.) of the logarithmic series for year 1995 (Mud Lake and Opax Mountain), spring samples  65  Figure 17: Alpha index values (+/- s.e.) of the logarithmic series for year 1996 (Mud Lake and Opax Mountain), spring samples  65  Figure 18: Alpha index values (+/- s.e.) of the logarithmic series for year 1997 (Mud Lake and Opax Mountain), spring samples  66  Figure 19: Shannon-Wiener diversity index values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples  70  F i g u r e 20: Simpson's diversity index values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples  70  Figure 21: Shannon-Wiener diversity index values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples  71  Figure 22: Simpson's diversity index values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples  71  Figure 23: Shannon-Wiener diversity index values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples  72  Figure 24: Shannon-Wiener diversity index values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples  72  Figure 25: Taxonomic distinctness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples  75  Figure 26: Taxonomic distinctness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples  75  Figure 27: Taxonomic distinctness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples  76  Figure 28: Average taxonomic distinctness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples  78  Figure 29: Average taxonomic distinctness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples  78  x  Figure 30: Average taxonomic distinctness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples Figure 31: Variation in taxonomic distinctness (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples Figure 32: Variation in taxonomic distinctness (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples Figure 33: Variation in taxonomic distinctness (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples Figure 34: The 95% 'probability funnel' for average taxonomic distinctness, 1995 spring samples (Mud Lake and Opax Mountain) Figure 35: The 95% 'probability funnel' for variation in taxonomic distinctness, 1995 spring samples (Mud Lake and Opax Mountain) Figure 36: The 95% 'probability funnel' for average taxonomic distinctness, 1996 spring samples (Mud Lake and Opax Mountain) Figure 37: The 95% 'probability funnel' for variation in taxonomic distinctness, 1996 spring samples (Mud Lake and Opax Mountain) Figure 38: The 95% 'probability funnel' for average taxonomic distinctness, 1997 spring samples (Mud Lake and Opax Mountain) Figure 39: The 95% 'probability funnel' for variation in taxonomic distinctness, 1997 spring samples (Mud Lake and Opax Mountain) Figure 40: Dendrogram for. hierarchical clustering of treatment data from Mud Lake and Opax Mountain (1995), using group-average linking of Bray-Curtis similarities calculated on 4 root-transformed abundance data t h  Figure 41: M D S of species abundances on treatment data for Mud Lake and Opax Mountain (1995) Figure 42: Dendrogram for hierarchical clustering of treatment data from Mud Lake and Opax Mountain (1996), using group-average linking of Bray-Curtis similarities calculated on 4 root-transformed abundance data t h  Figure 43: M D S of species abundances on treatment data for Mud Lake and Opax Mountain (1996)  xi  Figure 44: Dendrogram for hierarchical clustering of treatment data from Mud Lake and Opax Mountain (1997), using group-average linking of Bray-Curtis similarities calculated on 4 root-transformed abundance data  92  Figure 45: M D S of species abundances on treatment data for Mud Lake and Opax Mountain (1997)  92  Figure 46: Mean specimen number (+/- s.e.) of P. neobrunneus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  102  Figure 47: Mean specimen number (+/- s.e.) of C. taedatus agassii for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  102  Figure 48: Mean specimen number (+/- s.e.) of C. advena for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  103  Figure 49: Mean specimen number (+/- s.e.) of C. ingratus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  103  Figure 50: Mean specimen number (+/- s.e.) of P. adstrictus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  104  Figure 51: Mean specimen number (+/- s.e.) of S. marginatus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  104  Figure 52: Mean specimen number (+/- s.e.) of B. dyschirinum for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  105  Figure 53: Mean specimen number (+/- s.e.) of N. directus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  105  Figure 54: Mean specimen number (+/- s.e.) of C. unicolor for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  106  Figure 55: Mean specimen number (+/- s.e.) of S. americanus for seasons 1995-1997 (Mud Lake and Opax Mountain), based on spring samples only  106  th  xii  ACKNOWLEGMENTS -First of all I would like to thank my supervisor, Dr. Geoff Scudder, for his good advice, support, encouragement, and patience over all these years. I would also like to thank my two committee members, Dr. Judy Myers and Dr. Martin Adamson. -Next of all I would like to thank everybody who worked on the project, from the collectors and sorters in Kamloops and here, to all those who gave this neo-Luddite help with computers, and on how to write a thesis (Randy Daley, Dave Huggard, Diane Klimuk, Suzi Lavallee, Peggy Liu, Launi Lucas, Jocylyn McDowell, Karen Needham, Jonathan Parker, Robert Pollard, and of course the dozen or so monkeys whose random typing on my keyboard (with spellchecker) eventually produced this thesis). -Thanks also to Danny Schpeley of the Strickland Museum, University of Alberta, who verified all 58 species collected. - A very big thankyou to my parents (Don and Phyllis Jarrett) too, who supported me and encouraged me all this time. -Most of my thanks goes to my wife Michele though, who has been there for me always, and supported and helped me whenever she could. And also to my boy Samuel, who was always willing to help me with the ground beetles. -And I guess to all the 42,919 helpers out there too (you know who you are), who in taking that plunge, enabled me to study them closer, and to possibly help save a little piece of that IDF forest for their kids. I hope I have not disappointed you all.  This project supported financially by the Ministry of Forests, Forest Renewal British Columbia, and the Natural Sciences and Engineering Research Council of Canada.  xiii  INTRODUCTION  Forest Biodiversity Definition/Importance Biodiversity, as defined by the Convention on Biological Diversity at the United Nations Conference on Environment and Development in 1992, is "the variability among living organisms from all sources including terrestrial, marine and other aquatic ecosystems and ecological complexes of which they are apart: this includes diversity within species, between species and of ecosystems" (Heywood and Baste 1995). Not only does biodiversity encompass hierarchies of taxonomic and ecological scale, but it also encompasses temporal and geographic scales, as well as scaling in the body size of organisms (Stork 2003). To entomologists, however, biodiversity is insects, as insects comprise more than half of all described life forms, and possibly greater than 90% of all described and non-described life forms combined (Hawksworth and Kaiin-Arroyo 1995, Stork 2003). The interest in biodiversity lies not only in its variability, but also in what it produces (food, timber, industry products, pharmaceuticals, etc.), and maintains through ecosystem services (cleansing, recycling, and renewal) (Daily 1997). For example, in forest ecosystems biodiversity not only provides goods such as trees (for lumber and paper), but also game meat, fodder, and medicinal plants (McNeely 1994). Examples of ecosystem services that forests provide include: the stabilization of landscapes (Woodwell 1993), the protection of soils (Ehrlich and Ehrlich 1992), buffering against the spread of pests and disease (Woodwell 1995), the preservation of watershed functions (Burijnzeel 1990), the modulation of climate at both local and regional levels (Gash and Shuttleworth 1992,  1  Meher-Homji 1992), and helping to contain global warming through carbon stocks in both their plants and soils (Woodwell and Mackenzie 1995).  Biodiversity Numbers As mentioned above, insects represent the majority of life forms on the planet. Although about 850,000 to 1,000,000 insect species have been described, estimates of their true diversity range anywhere from 2-100 million species total, though a more likely number is probably around the 5-15 million range (Hawksworth and Kalin-Arroyo 1995, Stork 2003). Nevertheless, insects dominate the planet in terms of numbers of species. This is especially true in terrestrial ecosystems, where animal diversity has become synonymous with arthropod (insects and their allies) diversity (Asquith et al. 1990). Worldwide, natural forests are particularly rich in insect diversity. Although tropical forests cover only about 7% of the earth's land surface, they contain an estimated 50-90% of earth's species (Wilson 1988, Miller and Shores 1991). In Canada, temperate forests cover approximately 45% of the land mass (Canadian Forest Service 1994), but contain as much as two-thirds of the estimated 300,000 species of life forms in Canada (Boyle 1991). Estimates of the number of insect species in Canada come in at around 54,000 species (Danks 1979), and in British Columbia, the number is estimated to be around 35,000 species (Cannings and Cannings 1996), over 60% of the entire Canadian fauna!  Threats to Forest Biodiversity Unfortunately, there are numerous threats to biodiversity. These threats include both direct mechanisms (exploitation of wild living resources; expansion of agriculture,  2  forestry, and aquaculture; habitat loss and fragmentation; species introductions; pollution of soil, water, and atmosphere; and global climate change), and indirect mechanisms (human social organization; human population growth; natural resource consumption pattern; global trade; economic systems and polices that fail to value the environment and its resources; and inequity in the ownership, management and flow of benefits from both the use and conservation of biological resources) (Soule and Wilcox 1980, Diamond 1985, Pimm and Gilpin 1989). With forests containing the bulk of biodiversity, the loss of biodiversity in these habitats is consequently a major concern. In British Columbia, the loss of forest biodiversity is potentially linked to the following: general loss of forested habitat (through conversion to agriculture, suburbs, ski slopes), loss of specific forest structures (coarse woody debris, snags, large live trees), increased forest fragmentation, and the negative effects associated with too much forest edge (Fahrig and Merriam 1985, Andren and Angelstam 1988, Aubry et al. 1988, Hunter 1990, Franklin and Spies 1991, Morrison and Raphael 1993, Taylor and Fahrig 1993, O'Hara et al. 1994, Patten 1994, Dupuis etal. 1995, Bunnell and Chan-McLeod 1998). Although the number of forest species negatively affected by these trends is not completely known for British Columbia, if present trends continue, however, not only will species eventually disappear, but also will the important roles they play in ensuring a healthy forest ecosystem, and a sustainable forest industry.  Conservation of Biodiversity Worldwide public concern over the continued loss of species, and the destruction of ecosystems around the world, led to the Convention on Biological Diversity, which was  3  signed by 159 governments at the United Nations Conference on Environment and Development (UNCED) held in Brazil in 1992. The Convention builds on predecessors such as the World Conservation Strategy (IUCN, UNEP, W W F 1980), the (Brundtland) report of the World Commission on Environment and Development (Our Common Future) (WCED 1987), and Caring for the Earth: A Strategy for Sustainable Living (IUCN, UNEP, W W F 1991). Canada signed the Convention at U N C E D in 1992, and then became the first industrial country to ratify the Convention later that year. This was followed by the release of the Canadian Biodiversity Strategy in 1995, which put the Convention on Biological Diversity into a Canadian context. In the goal to maintain biological diversity in British Columbia's forested ecosystems, one approach has been to manage forests in a way as to mimic the natural disturbance regime, as is recommended by the Forest Practice Code (B.C. Ministry of Forests and B.C. Ministry of Environment, Lands and Parks 1995). The underlying assumption is that if forests are managed to resemble forests created by natural disturbance agents (insects, disease, fire, and wind), then all native species and ecological processes would naturally be maintained (B.C. Ministry of Forests and B.C. Ministry of Environment, Lands and Parks 1995). With this assumption in mind, biodiversity guidelines are recommended to address issues such as serai stages, patch size, connectivity, and stand structures (wildlife trees, coarse woody debris, tree species diversity, and understory vegetation diversity), in an effort to maintain forest biodiversity within the targeted social and economic constraints (B.C. Ministry of Forests and B.C. Ministry of Environment, Lands and Parks 1995). In this coarse filter ecosystem management approach, protecting habitat diversity, should in turn protect biodiversity. In addition to this, a number of  4  alternative forest management practices have been explored in recent years, in an effort to reduce the impact of forest harvesting on biodiversity. For example, besides clearcutting, methods such as patch cut, shelterwood, seed tree, group selection, and single tree selection have been explored (B.C. Ministry of Forests 1991, Kimmins 1997). However, there is little direct evidence to indicate the effectiveness of either the biodiversity guidelines, as suggested in the Forest Practice Code, or alternative harvesting methods, in reducing the impact on biodiversity in the different forested regions of British Columbia. This has subsequently led to the need for further research into this area.  O p a x M o u n t a i n Silvicultural Systems Research Project  The dry Douglas-fir forests of the Thompson and Nicola river basins have been managed by stand-level partial cutting for over 50 years (Klenner and Vyse 1998). Concern over the extensive use of 'continuous cover' silvicultural practices on issues such as regeneration, growth and yield, wildlife, pest management, and cattle grazing, led forest managers and other land managers in 1993 to form the Opax Mountain Silvicultural Systems Research Project (Vyse 1998). Initiated as a cooperative between the B C Ministry of Forests, University College of the Cariboo, Okanagan University College, and the University of British Columbia, its goal is to evaluate various alternative management practices, on the issues above, to the uniform partial cutting regimes of the past 50 years. To assess the effects that these various experimental harvesting approaches will have on the wildlife (biodiversity) at the Opax Mountain Silvicultural Systems Research  5  Project site, a range of indicator groups (e.g., ground dwelling arthropods, amphibians, shrews, mice and voles, sciurids, passerine birds, cavity nesting birds, and small carnivores and ungulates) were chosen that were likely to respond to the habitat structures and patterns being modified (Klenner and Huggard 1998a). In April of 1997, a workshop entitled "Managing the Dry Douglas-fir Forests of the Southern Interior" attempted to address the effects these experimental harvesting approaches had on the indicator groups chosen. A general overview of the results from these studies showed that not only do different indicators groups respond differently to the various experimental treatments, but individual species within the indicator groups respond differently as well (Huggard and Klenner 1998a, 1998b, Ferguson 1998, Klenner 1998a, 1998b, Klenner and Huggard 1998b). This thesis research focussed on investigating the effects the various harvesting approaches had on the ground dwelling arthropod indicator group, or more specifically, the Carabidae.  Carabidae  Diversity, distribution, biology and ecology Carabidae, or ground beetles, belong to the suborder Adephaga of the order Coleoptera. The terrestrial Adephaga, or Geadephaga, are represented primarily by the Carabidae (Thiele 1977). The main characteristics of carabids are: 1) the three basal visible sterna of the abdomen are coalescent, immobile, and often with more or less obliterated sutures; 2) the hind coxae are large and flat, fused to the metasternum and reach far beyond the posterior margin of the first visible (morphologically the second) sterna; 3) the number of sterna is six, only increasing to eight in the Brachinini; 4) the antennae are filiform (rarely  6  moniliform) and 11-jointed; and 5) the legs always have five tarsomeres (Snodgrass 1960, Lindroth 1961, Thiele 1977). The family Carabidae is one of the largest families of all insects, in terms of numbers of species, with approximately 40,000 species known worldwide (Erwin 1991). If one out of every five described organisms is a beetle (Evans and Bellamy 1996), then approximately one out of every 50 described organisms will be a ground beetle. In fact, there are only slighty fewer species of ground beetles than all of the described chordata (fish, birds, mammals, etc.) combined (Hawksworth and Kalin-Arroyo 1995). With such a diverse and successful fauna, I'm sure that if the British scientist J.B.S. Haldane was questioned further, he might have gone on to say that not only did the Creator have an "inordinate fondness for beetles" (Evans and Bellamy 1996), but that the ground beetles must have been one of his favorite creations. For example, in North America alone, 2636 species of carabids have been described, with 932 from Canada, and 505 from British Columbia (Bousquet and Larochelle 1993, Kavanaugh et al. 1998, Jarrett and Scudder 2001). Although the publications of Lindroth (1961-1969) and Wallis (1961) have greatly increased our taxonomic knowledge of Canadian carabid species, new species are still being described as recently as 1998 from B C (Kavanaugh et al. 1998). Thus, judging from their great diversity and abundance, it would seem to reason that carabids must play an important role in the nutritional chain of ecosystems (Thiele 1977), as well as in the regulation of phytophagous species numbers (Ball 1979). Carabids live on all continents, except Antarctica, and on most islands. Their geographic distribution ranges from well above the Arctic Circle to Tierra del Fuego and South Georgia in the Southern Hemisphere (Bousquet and Larochelle 1993).  7  Altitudinally, the range of carabids extends from sea level (and below) to 5300m in the Himalaya (Mani 1968). Carabids make their living in a wide variety of habitats, from cavernicolous to arboreal environments. Although no cavemicolous species are found in Canada (Lindroth 1961), a handful of species are known to climb trees in search of prey. Carabid species that inhabit the ground (geophiles or terracoles) are: hygrophiles, occupying riparian zones, sunlit marshes, and dark swamp forests; mesophiles, living in damp or wet forest or meadows, but independent of permanent surface water; and xerophiles, living in dry forest to grassland and desert situations (Ball and Bousquet 2001). In the tropics, over 60% of some regional carabid faunas are arboreal (Erwin 2001), making their living on tree trunks, branches, and some even on leaves (Ball and Bousquet 2001). Most temperate carabid species are univoltine, breeding in either spring or fall (Ball 1979). Females usually lay eggs singly, either on the ground, or in specifically constructed egg cells made of mud, twigs, or leaves (Dicker 1951, Forsythe 1987). The larvae of most carabid species have three instars, with some genera having two (Amaru and Harpalus), and others five (Brachinus) (Forsythe 1987). Pupation usually occurs in the ground. Autumn breeders will usually overwinter as larvae, while spring breeders will overwinter in the adult stage, either under the bark of dead trees, in hollow stems, at the base of grass tussocks, or in excavated chambers in the soil (Forsythe 1987). Many species of ground beetles also undergo a resting period, or diapause, in the larval or adult stage or, very rarely, as an egg (Forsythe 1987). Most species are also night active, or nocturnal (Ball and Bousquet 2001). Although the life cycle of most species is one year,  8  adults of the larger species tend to live for several years (e.g., Calosoma and Carabus) (Ball 1979). Most adult carabid beetles are polyphagous olfactory-tactile predators or scavengers (Brandmayr and Brandmayr 1980), eating dead or dying arthropods, or specializing on seeking active prey, such as mollusks, millipedes, or ants (Ball and Bousquet 2001). Some of the day-active optical predator species, such as the cicindelids, notiophilines, and loricerines, use primarily eyesight to capture active prey, including collembolans and other small soil-inhabiting arthropods (Ball and Bousquet 2001). Many other carabid species are phytophagous, eating seeds that have fallen to the ground, or which the beetles obtained in situ, on the plants (e.g., Amara and Harpalus). Some species are myrmecophiles and some are parasitoids, with the carabid larvae consuming its host over a period of days (Ball and Bousquet 2001). Carabids run along the ground, climb seldom, and either are totally incapable of flight (brachypterous species), or fly spontaneously on rare occasions only (Thiele 1977). However, only among the Cicindelinae and Bembidion is the power of flight regularly used for hunt and escape (Lindroth 1961). In most other species possessing functional wings, flight fulfils the purpose of migration between summer and winter habitats (Lindroth 1961).  Endangered carabids The IUCN Red List of Threatened Species lists seven species of carabid worldwide as being either near threatened, vulnerable, critically endangered, endangered, or extinct (IUCN 2002). The United States Endangered Species Act lists sixteen beetles in total,  9  seven of which are carabids that are either threatened or endangered in the U S A (U.S. Fish and Wildlife Service 2003). Canada's listing agency for endangered species, COSEWIC (Committee on the Status of Endangered Wildlife in Canada), only considers butterflies for listing in the insects to date (Cannings et al. 2001). In British Columbia, however, the British Columbia Conservation Data Centre (CDC) has assigned Red and Blue lists for tiger beetles (Cannings et al. 2001). In addition to this, Scudder's systematic list of potentially rare and endangered invertebrates for B C lists one species of carabid as being at risk, with thirteen others of special interest (Scudder 1994). Although only one recorded species of carabid has gone extinct since the year 1600 (IUCN 2002), this is certainly an underestimate of the true total of carabid extinctions, particularly with regard to the tropical fauna. Carabids become endangered when their habitats are either lost or degraded. Forest practices not only cause a loss of habitat for forest carabids, but they can also result in forests becoming overly fragmented. Fragmented forests negatively affect forest carabids by isolating populations, as well as increasing the amount of forest edge (Spence et al. 1996). Because isolated forest fragments contain fewer individuals of forest carabid species, they pose a greater risk for local extinctions due to stochastic population fluctuations, than do contiguous forest carabid populations (Niemela et al. 1993b). Furthermore, populations of the larger-bodied flightless carabid species in forest fragments cannot be maintained by dispersal, as distances between fragments become too great (Nield 1990, den Boer 1990, de Vries and den Boer 1990, Spence et al. 1997). The increased amount of edge caused by fragmentation also negatively impacts forest species by altering microclimates along edges, favoring the invasion of open habitat  10  species and habitat generalists (Halme and Niemela 1993, Spence et al. 1996). This not only results in increased competition along forest ecotones, but it also reduces the amount of forest interior habitat, critical for sensitive forest carabid species (Spence et al. 1996). This so called "edge effect" on forest fragments will ultimately result in edges having different species compositions and abundances, compared to uncut contiguous forests. In addition to the negative landscape effects caused by logging, forestry operations can also negatively impact species associated with certain micro-habitats, such as coarse woody debris, snags, large deciduous trees, and patches of wet swamp forest, that are characteristic of natural old growth forests (Berg et al. 1994, 1995, Haila 1994, Samuelsson et al. 1994, Siitonen 1994, Okland et al. 1996). Loss of these micro-habitat features will inevitably endanger those carabid species dependent on them. Futhermore, the effective suppression of forest fires adversely effects many pyrophilous species that require burnt substrates, and the post-fire successional stages that follow (Holliday 1984, Muona and Rutanen 1994). If the landscape and habitat changes caused by logging persist, they will eventually result in the homogenization of forested habitats, causing not only the decline of sensitive species associated with a heterogeneous forest habitat, but possibly even the more abundant forest generalist species in the long term (Niemela et al. 1993b). This homogenization of forests through logging will ultimately result in regional carabid diversity being depressed over the long term.  11  Indicators and Indices If one could measure every feature of a community or forest ecosystem, one could automatically determine the status and trend of any kind of biodiversity one cared about (Simberloff 1998). However, because of the size and complexity of forest ecosystems, this task would be impossible. Instead, indicator species, or groups of species, are used. Indicators species can: 1) reflect the biotic or abiotic state of the environment; 2) reveal evidence for, or the impact of, environmental change; and 3) indicate the diversity of other species, taxa, or entire communities within an area (Lawton and Gaston 2001). Because carabids are diverse, relatively well known taxonomically, have largely differentiated requirements, and respond rapidly to changed conditions in the environment, they have become a popular indicator group for predicting and assessing the effects of forest management, and other human disturbances to the environment (Frietag etal. 1973, Thiele 1977, Refseth 1980, Niemela etal. 1988, Honek 1988, Nield 1990, Szyszko 1990, Niemela et al. 1990b, 1992, Pearson and Cassola 1992, Niemela et al. 1993b). However, although the practice of monitoring carabids can tell much about the state of the epigeal environment, and repeated monitoring can provide information about changes to that environment, the use of monitoring carabids to indicate the diversity of other organisms is still debatable (Lawton and Gaston 2001). Also, because no single taxon reflects an entire habitat or ecosystem (Ricketts et al. 1999), ground beetles by themselves will never be sufficient indicators of the state, or change to the entire environment. Nevertheless, carabids have enormous potential as indicators in various biodiversity studies looking at the effects of habitat change on the ground dwelling arthropod guild.  12  In order to determine the state of, or change in the environment, however, one needs to use various diversity measures that look at changes in abundance, species richness, evenness, heterogeneity, distinctness, etc. in the indicator group chosen. In addition to using diversity measures for environmental assessment, diversity measures have also been used for conservation management purposes (Margurran 1988). In both cases, diversity measures can be used as an index of ecosystem wellbeing (Margurran 1988). However, diversity measures have their drawbacks. For instance, diversity indices, rankabundance curves, and multivariate analyses are all quantitative measures, with all species, taxonomically and functionally, implicitly treated as equal (Samways 1994). These measures also tell us nothing about the species composition. Phylogenetic diversity measures (e.g. taxonomic distinctness), nevertheless, can give an idea of the diversity of evolutionary pathways in the species sampled. Furthermore, diversity measures that measure the changes occurring in the environment fail to reveal the reasons as to why these changes are occurring (Samways 1994). This requires additional investigative work, with each organism's biology providing valuable clues (Lawton and Gaston 2001). Nevertheless, diversity measures will continue to be a useful tool for forest and other land managers, whether they are used for environmental monitoring purposes, or for direct conservation planning.  Opax carabids Carabids are used as ecological research subjects for a host of different reasons. Besides those already mentioned above, in regard to their use as indicator species, carabids are also: 1) an abundant and diverse family, occurring in most terrestrial  13  environments (Ball and Bousquet 2001); 2) are easily and inexpensively trapped i n large numbers for statistical study (Winchester and Scudder 1993, Marshall et al. 1994, K o i v u l a et al. 1999); and 3) being an epigeal group (in temperate regions), are appropriate subjects for studies comparing forested and logged treatments (Craig 1995). The carabids at the Opax Mountain study site were studied for the above reasons, as well as to compare to similar studies conducted recently in different biogeoclimatic zones across British Columbia (Craig 1995, Lemieux 1998, M c D o w e l l 1998, Lavallee 1999). It has long been known that carabids, for the most part, fall into different habitat association groups (Lindroth 1961, 1963, 1966, 1968, 1969, Thiele 1977). In temperate forest regions, species have been categorized as either forest specialists, forest generalists, habitat generalists, or open habitat species, depending on their preferences and adaptiveness to the local forest conditions (Niemela et al. 1992, Halme and Niemela 1993, Niemela et al. 1993a, 1993b). W h i l e a 100% adherence to any particular forest type represents an ideal not encountered in nature, the isolated occurrence of the odd carabid elsewhere, does not reduce the validity of grouping different species into their representative habitat association group (Thiele 1977). The work at Opax Mountain w i l l add to this baseline knowledge, by further categorizing the species inhabiting these I D F forests to their various habitat, or treatment groups. Previous researchers on carabid beetles i n forested environments have also investigated how carabids respond to coarse woody debris manipulation ( M c D o w e l l 1998), shrub removal (Parmenter and M a c M a h o n 1984, M c D o w e l l 1998), clearcutting (Sustek 1981, 1984, Lenski 1982b, Jennings et al. 1986, Langor et al. 1991, Lavallee 1999), forest "edge effects" (Spence etal. 1996, Niemela 1997, Lavallee 1999), scarification (Parry  14  and Roger 1986), climate change (Elias 1991), land reclamation (Day and Carthy 1988), forest fires (Holiday 1991, 1992), patch retention (Lemieux 1998), selective logging (McDowell 1998), and forest buffers along streams (Richardson et al. 2003). Conclusions based on these studies have provided forest managers with valuable information on how a successful forest epigeal invertebrate group has been impacted by various forest operations. The work at Opax Mountain adds to these previous studies, in that it provides managers with information on how carabids respond to not only different harvesting methods (individual-tree selection vs. patch cuts), but also to different percent levels of timber removal (20% vs. 50%), as well as to the presence of uncut reserves in 50% Individual-tree selection treatments. In accomplishing this, three unique indices will be used on the data from trap catches, namely taxonomic distinctness measures, multidimensional scaling graphs, and dominance structure tables. Another unique feature of this study is that it was carried out in the IDF biogeoclimatic zone of British Columbia, and in two different subzones within that biogeoclimatic zone. Thus, not only did this study yield new knowledge on species occurrence and range distribution for British Columbia (Jarrett and Scudder 2001), but it also increased our knowledge of rare carabid species, that might possibly be at risk of endangerment or extinction in British Columbia.  15  Objectives and Hypotheses  In examining the responses of both ground beetle assemblages and particular species to the various alternative harvesting practices at the Opax study site, the following objectives were developed: 1. To obtain an inventory of the epigeal carabid beetles occurring in the EDFdk and IDFxh forests of south-central British Columbia. 2. To assess how various alternative forestry practices impact both the diversity of ground beetle assemblages and abundance of specific sensitive species. 3. To impart information to wildlife/forest managers, on how to preserve carabid diversity in these human altered forest landscapes.  The hypotheses for this study are the following: 1. Species richness, evenness, and heterogeneity values will increase, while distinctness values decrease, as percent removal of timber increases from the research blocks (control - 20% - 35% - 50% removal). 2.  Species richness, evenness, and heterogeneity values will increase, while distinctness values decrease, as you move from control to individual-tree selection to patch-cut methods of harvest.  3. In multivariate analysis, treatment blocks harvested in a similar manner will cluster and map closer together, than treatment blocks where a similar percentage of forest was removed.  16  4. Dominance structure of the top ten species will deviate most from control block structure in treatments harvested by patch-cut methods as well as those treatments with a higher percentage removal of timber. 5. Intraspecific analysis will show that some carabid species will be caught in greater numbers in the higher percent forest removal treatments, others will be caught in fewer numbers in the higher percent forest removal treatments, some species will be caught in relatively similar numbers across all treatments, while other species will be caught almost exclusively in patch-cut treatments.  17  MATERIALS AND METHODS Site location and Description The Opax Mountain Silvicultural Systems Project site is located in the Interior Douglas-fir biogeoclimatic zone (Figure 1) southwest of McQueen Lake and Pass Lake about 20km northwest of Kamloops (Klenner and Vyse 1998). The study site consists of two areas, the Opax Mountain site and the Mud Lake site. The upper elevation site of Opax Mountain is located in the Interior Douglas-fir Dry Cool (IDFdkl) biogeoclimatic subzone, and covers the southerly slope of Opax Mountain at 1200-1370m elevation. The lower elevation site of Mud Lake is located in the Interior Douglas-fir Very Dry (IDFxh2) biogeoclimatic subzone, and surrounds Mud Lake at an elevation of 9501100m (Klenner and Vyse 1998).  Figure 1: Location of the Interior Douglas-fir zone in British Columbia (shaded area) and the location of the Opax Mountain Research Project (dark circle).  18  The Interior Douglas-fir (IDF) biogeoclimatic zone dominates the low to mid elevation landscape of south central British Columbia, between 49 degrees and 52 degrees 30'N latitude (Hope et al. 1991). Typically, the IDF occurs at elevations below the Montane Spruce biogeoclimatic zone and, where the valleys are deep enough, above the Ponderosae Pine biogeoclimatic zone (Hope et al. 1991). However, at the Opax Mountain Project study site, the IDF is bordered at lower elevations by the Bunchgrass biogeoclimatic zone. The IDF climate is characterized by warm, dry summers, a fairly long growing season, and cold winters (Hope et al. 1991). Twenty to fifty percent of the precipitation falls as snow, with substantial growing season moisture deficits common, and frosts occurring at anytime (Hope et al. 1991). There are seven subzones recognized in the IDF biogeoclimatic zone (Hope et al. 1991). The Mud Lake IDFxh2 subzone is characterized by an open and patchy spatial structure, with mainly Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco.), some spruce (Picea engelmannii Parry x glauca (Moench) Voss), lodgepole pine (Pinus contorta Dougl.), trembling aspen (Populus tremuloides Michx.), and paper birch (Betula papyrifera Marsh.) (Bealle-Statland 1998). The subdominant and intermediate trees have weak crowns and some top dieback, from a combination of poor site quality, drought, insect attack (particularly western spruce budworm {Choristoneura occidentalis Freeman}) and possibly lamenated root rot (Phellinus weirii (Murr.) Gilbertson) (BealleStatland 1998). Many of the largest diameter Douglas-fir trees have bore holes and pitch tubes from bark beetle (Dendroctonus pseudotsugae Hopkins) attack (Bealle-Statland 1998). The Opax Mountain IDFdkl subzone also has an open and patchy spatial structure, of approximately two-thirds Douglas-fir and one quarter to one third lodgepole  19  pine, with minor components of spruce and aspen (Bealle-Statland 1998). Growth is more vigorous and regeneration healthier than in the Mud Lake site, although many shaded understory trees have weak crowns and poor form (Bealle-Statland 1998). Less spruce budworm damage and top dieback is apparent. However, the lodgepole pine suffers from pine needle cast (Lophodermella concolor (Dearn.) Darker) disease and the large Douglas-fir have some beetle attack symptoms (Bealle-Statland 1998). Both sites have a herb-rich understory dominated by pinegrass (Calamagrostic rubescens Buckl), however, common snowberry (Symphoricarpos albus (L.) Blake) and ponderosa pine (Pinus ponderosa Dougl.) are common in the Mud Lake site, while lodgepole pine, soopolallie (Sheperdia canadensis (L.) Nutt.) and to a lesser extent kinnikkinnick (Arctostaphylos uva-ursi (L.) Spreng.) are more common at the Opax site (Miege et al. 1998). Eutric and Dystric Brunisols soils occur at both sites, as well as Gray Luvisols (Hope et al. 1991). The topography varies from steep slopes to depressions, and soil moisture ranges from submesic to hygric/subhygric (Kaipainen et al. 1998). The Mud Lake site was partially harvested in 1956 and 1957, probably with a diameterlimit method of single tree selection, while in the Opax Mountain site, some harvesting took place in 1957, but large areas remained uncut (Bealle-Statland 1998). Forestry is one of several important resource uses in the EDF, with cattle grazing, fur harvesting, and recreation also occurring in the IDF zone (Hope et al. 1991).  20  Logging Manipulations  The Opax Mountain Silvicultural Systems Project consists of two replicates of each of the following treatments in a randomized block design, at both Mud Lake and Opax Mountain (Figure 2): 20% merchantable volume removal using individual-tree selection (20% I.T.S.) (Units F, L). 50% merchantable volume removal using individual-tree selection (50% I.T.S.) (Units B , G). 35% merchantable volume removal, consisting of 70% of the treatment unit area harvested as 50% merchantable volume removal using individual-tree selection, and 30% of the treatment unit area retained as uncut reserves, around snags, large veterans, and broadleaf groups (50% I.T.S.(R)) (Units A , H). -  Patch cuts of 0.1, 0.4, and 1.6 ha on 20% of the treatment unit area (20% P.C.) (Units C, K ) .  -  Patch cuts of 0.1, 0.4, and 1.6 ha on 50% of the treatment unit area (50% P.C.) (Units E, J). Uncut controls (Units D, I).  Each replicate block is 20-25 ha, and harvesting occurred in winter 1993/1994. A range of site preparation methods were also applied to the site, but on a very small scale. Coarse woody debris was also experimentally manipulated on three 1 ha areas in each of the 50% patch cut treatments, and on the uncut controls, to provide sites with low, medium, and high coarse woody debris levels (Klenner and Vyse 1998).  21  20% volume removed 50% volume removed Uncut forest Patch cuts Roads  Harvested treatments at the Opax Mountain research site: uncut controls (units D, I); 20% I.T.S. (units F, L); 50% I.T.S. (units B, G); 20% P.C. (units C, K); 50% P.C. (units E, J); and 50% I.T.S.(R) (units A , H). F i g u r e 2:  S a m p l i n g Design  A triple-nested sampling design was used at the Opax Mountain Project research site (Huggard and Klenner 1998a). Five individual pitfall cups were set (according to RIC standards) in a 4m-radius circle to form a "trap circle" (Figure 3). Five trap circles, 35m apart in a cross pattern, formed a "set". Two sets of five circles each were located in each study block, with the sets at least 200m apart. In the winter, one chimney was used in the center of each circle of five summer traps. Trapping sessions were conducted in the autumn of 1993 (pre-treatment); the summer of 1994 (post-treatment); spring, summer, and autumn of 1995 and 1996; the spring of 1997; and in three post-treatment winters (Huggard and Klenner 1998a).  22  Figure 3: Pitfall trap sampling design at the Opax Mountain research site: (a) individual pitfall cup; (b) trap circle of five cups; (c) two sets of five circles per block.  Arthropod Sampling Technique Arthropods were trapped using arrays of small pitfall traps (400 ml plastic cups with opening diameter of 9.5 cm) set flush with the ground surface. A 30 x 30-cm board was held 15 cm above the pitfall trap on three stakes to keep rain and debris out of the trap. To set the trap, 100ml of 66% propylene glycol (a non-toxic and non-volatile liquid) was poured into the cup. Contents of the trap were collected 14 days after being set. A trapping "session" consisted of two consecutive 14-day collection periods. In winter, 1.3m chimneys of either plywood or P V C piping were held above the traps on raised cover boards. Winter traps were set through the chimneys, and collected two months later without disturbing the snow cover (Huggard and Klenner 1998a). Although pitfall catches are known to reflect the activity and density of carabids (Thiele 1977, Luff 1982, 1986), they are also known to be influenced by a number of other factors, including temperature and moisture (Ericson 1979, Honek 1988), surrounding vegetation (Greenslade 1964), material used for construction of the trap or as preservative (Luff 1975, Wagge 1985), and number (Obrtel 1971, Niemela et al. 1986), size, shape, and  23  arrangement of traps (Adis 1979). Despite these limitations, however, pitfall traps remain the best method for carrying out large-scale carabid studies.  Sorting a n d Identification  A total of 9,360 samples were processed from the following trapping sessions: autumn 1993 (pre-treatment), summer of 1994 (post-treatment), spring, summer and autumn of 1995 and 1996 (post-treatment), and the spring of 1997 (post-treatment). However, owing to the high number of traps, only two of the five traps per ring were processed in the autumn of 1995 and 1996. Arthropods from these samples were then sorted to order, and family where possible, with carabid beetles separated out and identified down to species. Identification of species was done using keys by Lindroth (1961, 1963, 1966, 1968, 1969) and Wallis (1961). Representatives of each species were pinned, with the remaining specimens stored in 7 and 20ml scintillation vials filled with 70% ethanol. Danny Schpeley (University of Alberta) kindly verified all pinned representatives. The reference collection at the Spencer Entomological Museum was also used in confirming identifications. Voucher material was prepared and placed in the Spencer Entomological Museum at U B C , the Royal British Columbia Museum and the Pacific Forestry Centre in Victoria, the Strickland Entomological Museum in Edmonton, the Canadian National Collection in Ottawa, the Oregon State University Arthropod Collection in Corvallis, Oregon, and the California Academy of Science, in San Francisco, California. Ecological data from Tables 1-6 (body length, geographical distribution, wing morphology, habitat association, and overwintering stage) was taken from Lindroth  24  (1961, 1963, 1966, 1968, 1969), Bousquet and Larochelle (1993), and Larochelle and Lariviere (2003).  Data Analysis After all samples were sorted, and carabids identified, data was entered into Microsoft Excel. Richness, evenness, heterogeneity, taxonomic distinctness, and multivariate analysis were then calculated using PRIMER Version 5.1.0. for windows. A l l other statistical calculations were done using SAS Version 8.0, and all figures made using Sigmaplot Version 4.0. Since total catches of carabids were highest for every year between the periods of late April to mid June, as well as numbers of species caught, the data analysis for this study (richness, evenness, heterogeneity, distinctness, multivariate and species analysis) focused strictly on the spring samples. Initial analysis of the data also revealed major differences between the Mud Lake and the Opax Mountain study site, both in terms of numbers of individual beetles collected, as well as total species caught. Because of these apparent differences, the two study sites were analyzed separately for this study. However, this unfortunately left only two replicates per study site, with five trap circles each.  Species Richness Species richness, as coined by Mcintosh (1967), is meant to describe the number of species in a community. Although the oldest and most simple measure of diversity,  25  species richness measures are problematic when used to describe communities based on different sample sizes. This problem was overcome by Sanders (1968) who proposed the method of Rarefaction, which is a statistical method for estimating the number of species expected in a random sample, if all samples were reduced to a standard size. Because carabids show a non-random spatial distribution both within and between habitats (Thiele 1977, Luff 1986, Niemela 1990, Niemela and Halme 1992, Niemela et al. 1992) overestimating species richness using the method of Rarefaction is a possibility (Fager 1972). To reduce this bias, a large number of traps were spread over the two study sites, as is recommended by Krebs (1989).  Evenness As is common in many plant and animal communities, with carabids being no exception, few species are very abundant, some have a medium abundance, while most are represented by only a few individuals (Magurran 1988, Niemela et al. 1990a). Because of this characteristic pattern of species abundance, ecologists have devised evenness measures that attempt to quantify this unequal representation against a hypothetical community in which all species are equally common (Krebs 1989). Pielou's J' evenness measure, which is based on the Shannon-Wiener function (Peet 1974), was one such measure that was calculated for this study. Equitability or evenness can also be observed graphically in terms of the underlying species abundance distribution. This was done using Whittaker plots (1965), with percent abundance plotted logarithmically on the Y-axis, against species rank on the X-axis.  26  Heterogeneity The term heterogeneity, as first applied by Good (1953), incorporates both the number of species (species richness) and the distribution of individuals among those species (evenness) into a single value (Pielou 1975, Ludwig and Reynolds 1988). The measurement of diversity by means of heterogeneity indices, however, has proceeded along two different paths (Krebs 1989). One approach is to use the statistical sampling theory to investigate how communities are structured, while the other approach looks to information theory as an appropriate measure of diversity (Krebs 1989). An example of the first approach, which was calculated for this study, is the alpha diversity index from the logarithmic (or log) series species abundance distribution. Even when the logarithmic series distribution is not the best descriptor of the data, the alpha index is still a favorable index of diversity (Taylor 1978, Kempton and Taylor 1974, 1976). The second approach in measuring diversity is to use non-parametric measures of heterogeneity, which make no assumptions about the shape of species abundance curves (Krebs 1989). The two nonparametric heterogeneity measures calculated in this study are the Shannon-Wiener (H') and Simpson's (1/D') diversity indices. The Shannon-Wiener index measures the average degree of 'uncertainty' in predicting to what species an individual chosen at random from a collection of S species and N individuals will belong (Pielou 1975). The Simpson's index, on the other hand, measures the probability that two individuals selected at random from a sample will belong to the same species (Peet 1974). The Shannon-Wiener index is an example of a type 1 index, showing most sensitivity to changes in the rare species in the community, while the Simpson's index is an example of  27  a type 2 index, being more sensitive to changes in the most common species in the community (Peet 1974).  Taxonomic Distinctness The phylogenetic structure of an assemblage is an important biological attribute of animal communities. Although measurements incorporating phylogenetic structure have been used in selecting species or reserves of greatest conservation priority (Faith 1994, Humphries et al. 1995), measuring phylogenetic structure as a further measure of biodiversity, was first introduced by Warwick and Clarke (1995) with their concept of taxonomic distinctness. Taxonomic distinctness indices can measure the average distance apart of all pairs of individuals (taxonomic distinctness or TD) and species (average taxonomic distinctness or aveTD) traced through a taxonomic tree, as well as variability in structure across the tree (variation in taxonomic distinctness or varTD). In addition to calculating these three taxonomic distinctness measures, a 'probability funnel' (Clarke and Warwick 1998) was constructed based on the master list of carabids collected during the study, in order to test aveTD and varTD values in each of the different treatments for their departure from the 'expected' aveTD and varTD values.  Multivariate Analysis Multivariate methods are characterized by the fact that they base their comparisons of two or more samples on the extent to which these samples share particular species, at comparable levels of abundance (Clarke and Warwick 1994). The two multivariate techniques used in this study are hierarchical agglomerate clustering (Everitt 1980) and  28  non-metric multi-dimensional scaling, or simply M D S (Kruskal and Wish 1978). Both these techniques were based on 4 -root transformed Bray-Curtis similarity coefficients, th  which compute the similarity between every pair of samples, or treatments in this case. In the hierarchical agglomerate clustering technique, treatments are successfully fused into groups, starting with the highest mutual similarities, then gradually lowering the similarity level at which groups are formed, resulting in a dendrogram, or tree diagram (Clarke and Warwick 1994). In the M D S technique, treatments are usually 'mapped' in two-dimensions, in such a way that the rank order of the distances between treatments on a map exactly agrees with the rank order of the matching similarities, taken from the triangular similarity matrix (Clarke and Warwick 1994). When used in combination, these two techniques can be an effective way in analyzing similarities in species compositions between two or more treatments.  Dominance Structure The dominance structure of the carabid assemblages was also investigated in this study. At both Mud Lake and Opax Mountain research sites, I recorded the dominance position and percent abundance of the ten most abundant carabid species found in control blocks, and followed the dominance position of each of the ten species in the five harvested treatments, as well as their percent abundance. This was done for both spring and entire year samples at each site. B y investigating the dominance structure of carabid assemblages, a more complete picture can be made on how logging impacts carabid communities. Although only the  29  top ten species from control blocks were analyzed, they accounted for at least 90% of the total catch from each treatment.  Species Analysis Because rare species can vary considerably both in their presence and abundance, such species cannot have functional linkages central to the integrity of communities (Niemela and Spence 1994). For this reason, only the ten most abundant species (>200), collected from spring samples, were analyzed for their response to the six different treatments. Two replicates, of five trap circles each, resulted in a total of ten numbers representing "numbers per trap" for species abundance in each treatment for both study sites. The mean and standard error of these ten numbers were plotted as histograms for each treatment in each site, for the combined 1995-97 data.  Statistical Tests Since repeated estimates of diversity are usually normally distributed, analysis of variance tests (ANOVA) can be used to test for significant differences between the diversity of different sites (Magurran 1988). A single factor analysis of variance, or Ftest, was used in this study to test for treatment differences in species richness, evenness, alpha index, Shannon-Wiener index, Simpson's index, taxonomic distinctness, average taxonomic distinctness, and variation in taxonomic distinctness. If significant differences were detected between treatments, the Tukey multiple comparison test was used to determine which treatments differed significantly. The Kruskal-Wallis single-factor analysis of variance by rank was also used, in this case to test for treatment differences in  30  each of the abundant ten species. This test is recommended when samples do not come from a normal population and/or the variances are heterogeneous, as was the case for this study. If significant differences occurred between treatments, a non-parametric multiple comparison test, similar to the Tukey test, was carried out on the mean numbers of species. This calculation was done by hand, following the method outlined by Zar (1984).  31  RESULTS A total of 42,919 carabid beetles were collected from the Opax Mountain Silvicultural Systems Project site. From the complete set of 9,360 samples analyzed, spanning over 131,040 trap days (a trap day is a single trap operated over a 24-hour period), 58 species from 24 genera were identified, with two species, Amara aenea (DeGeer) and Badister obtusus LeConte new to B C (Jarrett and Scudder 2001). The two most abundant species, Carabus taedatus agassii LeConte and Pterostichus neobrunneus Lindroth account for almost 53% of the total number of beetles caught. Four other species were present in numbers greater than 1000, eleven species were represented by more than 100 specimens, and 30 species were represented by less than ten specimens each, including seventeen singleton species (species represented by one specimen) (Table 1). A total of 49 species, or roughly 85%, were either macropterous, dimorphic, or polymorphic with respect to wing morphology, while the remaining nine species were brachypterous. A total of 36 species, approximately 59%, overwinter in the adult stage, eleven species overwinter as larvae, ten species are known to either overwinter as adults or larvae, and one species is unresolved as to overwintering stage. Also, four alien species of carabids were collected, comprising a total of only five specimens.  M u d Lake site A total of 16,734 carabid beetles were collected during the complete sampling period at the Mud Lake site of the Opax Mountain Silvicultural Systems Project. Of these, 49 species from 22 genera were identified, with 22 species unique to the Mud Lake site.  32  Carabus taedatus agassii and Pterostichus neobrunneus were the two most abundant species, accounting for more than 46% of the total beetles caught. Two other species were present in numbers greater than 1000, ten species were represented by numbers greater than 100 individuals, and 22 species were represented by less than ten individuals each, including thirteen singleton species (Table 2). A total of 43 species were either macropterous, dimorphic, or polymorphic with respect to wing morphology, with the remaining six species brachypterous. A total of 30 species overwinter in the adult stage, eleven species overwinter as larvae, seven species are known to overwinter as adult or larvae, and one species is unresolved as to overwintering stage. Also, three alien species of carabids were collected from the Mud Lake site, comprising a total of four specimens. Results from spring samples at Mud Lake (Table 7) showed that 20% I.T.S. treatments contained the most individuals, with 50% I.T.S. treatments containing the least. For species number, richness, and Shannon-Wiener diversity index, 50% patch-cut treatments had the highest value, while control blocks had the lowest. For the Simpson's diversity index, however, 20% patch-cut treatments had the highest value, with 20% I.T.S. treatments recording the lowest. Compared to individual-tree selection treatments (20% and 50% I.T.S.), patch-cut treatments (20% and 50% P.C.) had higher values for species richness, Shannon-Wiener and Simpson's diversity indices, however, numbers of individuals and species were inconclusive. Compared to 20% removal treatments (20% I.T.S. and 20% P . C ) , 50% removal treatments (50% I.T.S. and 50% P.C.) recorded higher values for species number, richness, and Shannon-Wiener diversity index, with inconclusive results for the Simpson's diversity index. While 50% LT.S.(R) treatments  33  were intermediate in numbers of individuals, richness, Shannon-Wiener and Simpson's diversity indices, when compared to 20% and 50% I.T.S. treatments, they contained more species than both. 50% patch-cut treatments contained the highest number of immigrant species (new species found only after logging), with both 20% removal treatments containing the lowest number (Table 5). 20% patch-cut and 50% I.T.S. treatments contained the highest number of emigrant species (species disappearing after logging), with 50% LT.S.(R) treatments the lowest. Results from entire year samples at Mud Lake (Table 8) showed that, similar to spring samples (Table 7), 20% I.T.S. treatments contained the most individuals, with 50% I.T.S. treatments the fewest. For number of species, 50% I.T.S. treatments contained the greatest number, with control blocks the fewest. For species richness, Shannon-Wiener and Simpson's diversity indices, 50% patch-cut treatments had the greatest value, while control blocks (for richness and Shannon-Wiener index) and 50% I.T.S. treatments (for Simpson's index) had the least. Similar to spring samples (Table 7), patch-cut treatments had a greater richness, Shannon-Wiener and Simpson's diversity index value, when compared to individual-tree selection treatments, but showed inconclusive results for both species number and number of individuals. Compared to 20% removal treatments, 50% removal treatments had a higher richness, with other results inconclusive. While 50% I.T.S.(R) treatments were intermediate in numbers of individuals, species richness, and number of species, when compared to 20% and 50% I.T.S. treatments, they recorded higher values in both the Shannon-Wiener and Simpson's diversity indices. 50% I.T.S. treatments contained the highest number of immigrant species, with 20% I.T.S.  34  treatments the least (Table 2). The 20% I.T.S. treatments contained the higher number of emigrant species, with 50% I.T.S(R) treatments the fewest.  Opax Mountain site A total of 26,185 carabid beetles were collected during the complete sampling period at the Opax Mountain site of the Opax Mountain Silvicultural Systems Project. Of these, 36 species from 19 genera were identified, with nine species unique to the Opax Mountain site. Similar to the Mud Lake site,  Carabus  taedatus agassii  and Pterostichus  neobrunneus  were the two most abundant species, accounting for more than 57% of the total number of beetles caught. Three other species were present in numbers greater than 1000, six species were represented by more than 100 individuals, and fifteen species were represented by less than ten individuals, including seven singleton species (Table 3). A total of 28 species were either macropterous, dimorphic, or polymorphic with respect to wing morphology, with the remaining eight species brachypterous. A total of 23 species overwinter in the adult stage, six species overwinter as larvae, and seven species can overwinter as adults or larvae. One individual alien species was also collected from the Opax Mountain site. Results from spring samples at Opax Mountain (Table 9) showed that 50% I.T.S. treatments contained the greatest number of individuals, with 50% patch-cut treatments the fewest. For species number, 50% patch-cut treatments contained the highest number, with 20% I.T.S. treatments the lowest. For species richness, Shannon-Wiener and Simpson's diversity indices, 50% patch-cut treatments had the greatest value, with control  35  blocks (for Shannon-Wiener and Simpson's index) and 50% I.T.S. treatments (for species richness) the lowest. Compared to individual-tree selection treatments, patch-cut treatments contained the greater number of species and richness, with inconclusive results for the other indices. Compared to 20% removal treatments, 50% removal treatments had the greater number of species, with inconclusive results for the other indices. While 50% I.T.S.(R) treatments were intermediate in numbers of individuals, Shannon-Wiener and Simpson's diversity indices, compared to 20% and 50% I.T.S. treatments, they were greater in species number and richness than both. 50% patch-cut treatments had the highest number of immigrant species, with 20% I.T.S. treatments the lowest (Table 6). 50% I.T.S.(R) treatments had the highest number of emigrant species, while both patchcut treatments contained the lowest. Results from the entire year samples at Opax Mountain (Table 10) showed that, similar to spring samples (Table 9), 50% I.T.S. treatments contained the greatest number of individuals, with 50% patch-cut treatments the fewest. Both 50% patch-cut and 50% I.T.S.(R) treatments contained the greatest number of species, with 20% I.T.S. treatments the least. Similar to spring samples (Table 9), 50% patch-cut treatments had the highest richness, Shannon-Wiener and Simpson's diversity index values, with control blocks (for Shannon-Wiener and Simpson's indices) and 50% I.T.S. treatments (for richness) the lowest value. Compared to individual-tree selection treatments, patch-cut treatments had a greater species number, Shannon-Wiener and Simpson's index value, but showed inconclusive results for other indices. Compared to 20% removal treatments, 50% removal treatments were greater for species number and Simpson's diversity index values, but showed inconclusive results for the other indices. While 50% I.T.S.(R)  36  treatments were intermediate in number of individuals and the Simpson's diversity index value, when compared to 20% and 50% individual-tree selection treatments, they were greater than both for species number, richness, and the Shannon-Wiener index value. Both 50% patch-cut and 50% I.T.S.(R) treatments contained the greatest number of immigrant species, with 20% I.T.S. treatments the least (Table 3). 20% I.T.S. treatments surprisingly contained the higher number of emigrant species, with 20% patch-cut treatments the lowest.  37  Table 1: Total number of specimens caught in all experimental treatments at the Opax Mountain Silvicultural Systems Project site.  Trachypachus  holmbergi  Mannerheim  Pre-harvest  Control  Dist.  W.morph.  Habitat  O/W Stage  0  2  20%I.T.S. 20%P.C. 1  8  0  15  17  43  0.100  3.8 -5.8  N  Ma  O  A  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Total  % of Total Length(mm)  Leistus ferruginosus  Mannerheim  0  1  0  0  1  0  0  2  0.005  7.8 -9.3  N  Ma  R/F  L  Notiophilus  Casey  10  120  182  147  95  205  108  867  2.020  4.8 -5.7  N  D  O/F/R  A  directus  Loricera pilicornis Cicindela  pilicomis  Calosoma  1  0  0  0  0  0  0  1  0.002  7.0 -8.5  H  Ma  R/C  A  0  0  0  2  1  0  0  3  0.007  14.0 -16.0  N  Ma  G  L  (LeConte)  0  0  0  0  0  0  1  1  0.002  15.0 -20.0  N  B  O/M  L  Say  3  0  0  0  1  0  24  28  0.065  16.0 -24.0  N  D  O/F/M/C  A  225  2686  2376  1909  1841  2058  1993  13088  30.495  16.0 -23.0  N  B  O/F/M/G/C  A  1  0  1  0  0  0  1  3  0.007  16.0 -20.0  N  B  F  LI A  7  135  108  35  77  29  32  423  0.986  17.0 -24.0  N  B  F/R/C  L/A  71  360  398  234  156  226  274  1719  4.005  11.5 -19.0  N  B  F/R/C/T  L  3  7  7  8  1  8  0  34  0.079  6.2 -9.8  H  Ma  O/F/M  A  0  0  0  1  0  0  0  1  0.002  3.6 -4.1  I  D  O/M/C/F  L/A  wilkesii  Carabus serratus  (Fabricius)**  Casey  nebraskana  Carabus taedatus agassii Scaphinotus  relictus  Scaphinotus  angusticollis  Scaphinotus  marginatus  Miscodera  arctica  LeConte  (G.H. Horn) (Fischer von Waldheim) (Fischer von Waldheim)  (Paykull)  Trechus obtusus  Erichson  Trechus tenuiscapus  Lindroth**  Bembidion  dyschirinum  Bembidion  erasum  Bembidion  iridescens  Bembidion  grapii  Bembidion  rupicola  LeConte  LeConte (LeConte)  2  0  0  0  0  0  0  2  0.005  4.1 -4.9  N  B  F  A  87  22  33  75  21  155  44  437  1.018  2.9 -3.8  N  D  O/F  A  0  0  0  0  0  1  0  1  0.002  3.0 -4.2  N  Ma  R/F  A  0  0  0  6  0  0  0  6  0.014  3.8 -4.8  N  Ma  R/F  A  Gyllenhal  9  1  1  19  1  43  1  75  0.175  4.0 -4.8  H  Po  O/M/C/F  A  (Kirby)  0  0  1  0  0  1  0  2  0.005  4.1 -5.5  N  Ma  O/F/C/R/M  A  Chaudoir  0  0  0  0  0  0  1  1  0.002  8.3- 12.0  N  Ma  R/O  L/A  0  0  0  0  0  0  1  1  0.002  9.5- 12.5  H  D  T  A  383  303  341  495  527  903  795  3747  8.730  9.5- 13.0  H  Ma  G/O/F/C/R  L/A  17  4  8  92  31  2  93  247  0.576  12.0 -15.0  N  Ma  F  A  9  2  19  2  3  1  8  44  0.103  9.5- 12.0  N  D  F/R  A  0  0  0  0  0  0  1  1  0.002  12.0 -19.0  I  D  C/F/M/R/G  L  306  1667  1606  1783  1810  1207  1234  9613  22.398  11.5 -14.2  N  B  F  A  Patrobus stygicus Stereocerus  haematopus  Pterostichus  adstrictus  Eschscholtz  (Dejean)  Pterostichus  oregonus  LeConte  Pterostichus  pensylvanicus  Pterostichus  melanarius  Pterostichus  neobrunneus  Pterostichus  riparius  LeConte (llliger) Lindroth  (Dejean)  0  0  0  0  4  0  0  4  0.009  6.5 -8.0  N  B  R/F  A  (Kirby)  0  0  1  2  1  2  0  6  0.014  7.8- 10.8  N  Ma  O/F/C/M  L  Amaraobesa  (Say)  0  0  0  0  1  0  0  1  0.002  9.0- 12.5  N  D  O/R/C/M  L  Amaraaenea  (DeGeer)  0  0  0  0  0  0  1  1  0.002  6.2 -8.8  I  Ma  M/O/G/C  A  0  0  0  1  0  0  0  1  0.002  7.6- 10.0  N  Ma  ?  ?  Ma  O/M/G/C  A  Ma  O/G/M/F  A  Amaralatior  Amara conflata Amara ellipsis Amara erratica  LeConte (Casey) (Duftschmid)  0  0  0  10  18  42  3  73  0.193  7.0 -9.0  N  0  0  0  1  1  1  0  3  0.005  6.5 -8.7  H  Table 1: Total number of specimens caught in all experimental treatments at the Opax Mountain Silvicultural Systems Project site (cont.). Dist.  W.morph.  Habitat  O/W Stage  0  0  0  0  1  1  2  0.005  5.6 - 7.2  I  Ma  O/C/F/M  A  0  0  0  0  1  0.002  6.5 - 9.0  N  Ma  O/R/C/M  A  Pre-harvest Control 20%I.T.S. 20%P.C. 50%I.T.S. 50%P.C. 50%I.T.S.(R) Amara familiaris (Duftschmid)  0  Total  % of Total Length(mm)  Amara farcta LeConte  0  0  1  Amara idahoana (Casey)  5  6  5  17  13  54  9  109  0.254  5.0-6.5  N  D  O/M/G  L  Amara laevipennis Kirby  0  1  1  35  10  122  4  173  0.382  5.8-7.1  N  Ma  O/C/F/M/G  A  Amara littoralis Mannerheim  0  0  3  13  9  20  4  49  0.112  6.2 - 9.3  N  Ma  O/C/F/M/G  A  Amara lunicollis Schiodte  0  0  0  7  5  6  9  27  0.063  7.3 - 9.0  H  Ma  O/C/F/M/G  A  Badister obtusus LeConte**  1  0  0  0  0  0  0  1  0.002  5.1 -6.5  N  Ma  F/C  A  Trichocellus cognatus (Gyllenhal)  1  1  0  0  1  5  0  8  0.019  3.5 - 5.2  H  Ma  G/F/M/C  A  Harpalus animosus Casey  0  1  0  3  8  4  1  17  0.040  12.5 -14.0  N  Ma  G/M/O/R  L/A  Harpalus fulvilabris Mannerheim  0  0  1  0  0  0  0  1  0.002  8.8-10.8  N  D  O/F  A  Harpalus nigritarsus C R . Sahlberg  2  7  27  10  14  28  15  103  0.240  7.0 - 9.2  H  Ma  O/G/M  A  Harpalus innocuus LeConte  0  0  2  0  1  2  1  6  0.014  8.2 - 9.7  N  Ma  O/M  A  Harpalus plenalis Casey  0  0  0  0  0  0  1  1  0.002  7.5 - 8.4  N  Ma  0  L  Ma  O/F/G/M/C  L/A  Harpalus opacipennis (Haldeman)  0  0  1  4  3  12  6  26  0.061  7.5 - 9.3  N  Harpalus solitaris Dejean  0  1  0  0  0  0  0  1  0.002  8.8-10.4  H  Ma  O/F/M  L/A  Harpalus fuscipalpis Sturm  0  0  0  0  2  0  0  2  0.005  7.3 - 9.8  H  Ma  O/C/G/M  L/A  Calathus ingratus Dejean  87  719  1174  764  741  852  657  4994  11.636  7.0-11.1  N  D  Calathus advena (LeConte)  183  927  683  741  934  451  1249  5168  12.041  8.0-11.8  N  Ma  F/M/R  L  721  1.680  8.7-11.2  N  D  C/F  L  Synuchus impunctatus (Say)  15  182  100  139  43  132  110  O/  F/11  Gl C /M  L/A  Sericoda quadripunctala (DeGeer)  0  0  0  1  0  0  0  1  0.002  4.2 - 6.0  H  Ma  P/F  A  Agonum retractum LeConte  8  0  3  1  1  1  1  15  0.035  6.2 - 7.6  N  D  F  A  Agonum cupreum Dejean  0  0  0  0  0  1  0  1  0.002  7.0 - 9.5  N  D  O/G/M/C  A  Cymindis cribicollis Dejean  5  24  42  12  24  8  33  148  0.345  8.4-11.0  N  D  F/M/C  A  Cymindis unicolor Kirby  9  88  115  110  78  118  85  603  1.405  8.0 - 9.5  N  B  O/T/M  A  0.610  2.7 - 3.5  N  D  O/F/G/M/C  A  Syntomus americanus (Dejean)  7  8  28  66  20  93  40  262 42919  TOTAL  1457  7275  7269  6753  6498  6809  6858  % TOTAL  3.39  16.95  16.94  15.73  15.14  15.87  15.98  # SPECIES  26  25  30  34  36  35  36  58  #GENERA  17  15  14  17  16  15  16  24  # IMMIGRANT SPECIES (AFTER LOGGING)  9  12  13  12  15  30  # EMIGRANT SPECIES (AFTER LOGGING)  4  3  2  2  4  1  Note: N=nearctic, H=holarctic, ^introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, 0=open, R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, A=adult, L= larval, P=pyrophilous, W.morph=wing morphology, 0/W=overwintering stage, Dist.= distribution, I.T.S.=individual tree selection, P.C.=patch-cut, (R)=reserves, **=unique to pre-harvest.  Table 2: Total number of specimens caught in all experimental treatments at the Mud Lake site (List of species as in Table 1). 50%I.T.S.  50%P.C  50%I.T.S.(R)  Total  Dist.  W.morph.  0  2  1  3  0  13  16  35  0.209  3.8 - 5.8  N  Ma  O  Pre-harvest Control Trachypachus  holmbergi  Mannerheim  20%I.T.S. 2 0 % P . C  % of Total Length(mm)  . Habitat  O/W Stage A  Leistus ferruginosus  Mannerheim  0  0  0  0  0  0  0  0  0.000  7.8 - 9.3  N  Ma  R/F  L  Noliophilus  Casey  4  12  34  22  14  40  26  152  0.908  4.8 - 5.7  N  D  O/F/R  A  directus  'Loricera pilicomis 'Cicindela "Calosoma 'Carabus Carabus  pilicornis  wilkesii serratus  (Fabricius)"  1  0  0  0  0  0  0  1  0.006  7.0-8.5  H  Ma  R/C  A  Casey  0  0  0  2  1  0  0  3  0.018  14.0-16.0  N  Ma  G  L  (LeConte)  0  0  0  0  0  0  1  1  0.006  15.0-20.0  N  B  O/M  L  3  0  0  0  1  0  24  28  0.167  16.0-24.0  N  D  O/F/M/C  A  178  906  785  692  585  629  651  4426  26.449  16.0-23.0  N  B  O/F/M/G/C  A  nebraskana  Say  taedatus agassii  Scaphinotus  relictus  Scaphinotus  angusticollis  Scaphinotus  marginatus  Miscodera  LeConte  (G.H. Horn) (Fischer von Waldheim) (Fischer von Waldheim)  0  0  0  0  0  0  0  0  0.000  16.0-20.0  N  B  F  L/A  7  49  100  18  14  17  12  217  1.297  17.0-24.0  N  B  F/R/C  L/A  54  236  245  164  59  142  83  983  5.874  11.5-19.0  N  B  F/R/C/T  L  arctica  (Paykull)  0  0  2  0  0  0  0  2  0.012  6.2 - 9.8  H  Ma  O/F/M  A  "Trechus obtusus  Erichson  0  0  0  1  0  0  0  1  0.006  3.6-4.1  I  D  O/M/C/F  L/A  Trechus tenuiscapus  Lindroth  0  0  0  0  0  0  0  0  0.000  4.1 -4.9  N  B  F  A  11  5  11  42  12  57  16  154  0.920  2.9 - 3.8  N  D  O/F  A  0  0  0  0  0  0  0  0  0.000  3.0 - 4.2  N  Ma  R/F  A  0  0  0  6  0  0  0  6  0.036  3.8 - 4.8  N  Ma  R/F  A  Gyllenhal  4  0  0  6  0  11  0  21  0.125  4.0 - 4.8  H  Po  O/M/C/F  A  (Kirby)  0  0  1  0  0  0  0  1  0.006  4.1 -5.5  N  Ma  O/F/C/R/M  A  Chaudoir  0  0  0  0  0  0  0  0  0.000  8.3-12.0  N  Ma  R/O  L/A  Bembidion  dyschirinum  Bembidion  erasum  'Bembidion  LeConte  LeConte  iridescens  Bembidion  grapii  Bembidion  rupicola  Patrobus stygicus 'Stereocerus Pterostichus  (LeConte)  haematopus  (Dejean)  adstrictus  Eschscholtz  Pterostichus  oregonus  LeConte  Pterostichus  pensylvanicus  "Pterostichus  melanarius  Pterostichus  neobrunneus  Pterostichus  riparius  LeConte (llliger) Lindroth  0  0  0  0  0  0  1  1  0.006  9:5-12.5  H  D  T  A  336  236  244  290  192  630  612  2540  15.179  9.5-13.0  H  Ma  G/O/F/C/R  L/A  15  3  8  91  31  2  65  215  1.285  12.0-15.0  N  Ma  F  A  6  1  0  0  1  0  2  10  0.060  9.5-12.0  N  D  F/R  A  0  0  1  1  0.006  12.0-19.0  I  D  C/F/M/R/G  L  B  F  A  0  0  0  0  152  581  854  430  273  590  416  3296  19.696  11.5-14.2  N  0  0  0  0  0  0  0  0  0.000  6.5-8.0  N  B  R/F  A  Amara latior (Kirby)  0  0  1  2  1  0  0  4  0.024  7.8-10.8  N  Ma  O/F/C/M  L  'Amara obesa  0  0  0  0  1  0  0  1  0.006  9.0-12.5  N  D  O/R/C/M  L  0  0  0  0  0  0  0  0  0.000  6.2 - 8.8  I  Ma  M/O/G/C  A  0  0  0  1  0  0  0  1  0.006  7.6-10.0  N  Ma  ?  ?  0  0  0  7  18  26  0  51  0.305  7.0 - 9.0  N  Ma  O/M/G/C  A  0  0  0  1  1  1  0  3  0.018  6.5 - 8.7  H  Ma  O/G/M/F  A  Amara aenea  (Say) (DeGeer)  "Amara conflata Amara ellipsis "Amara erratica  (Dejean)  LeConte (Casey) (Duftschmid)  Table 2: Total number of specimens caught in all experimental treatments at the Mud Lake site (cont.). 50%I.T.S.  50%P.C.  50%I.T.S.(R)  Total  Dist.  W.morph.  Habitat  O/W Stage  0  0  0  0  0  1  1  2  0.012  5.6 - 7.2  I  Ma  O/C/F/M  A  0  0  1  0  0  0  0  1  0.006  6.5 - 9.0  N  Ma  O/R/C/M  A  4  3  3  14  7  9  4  44  0.263  5.0-6.5  N  D  O/M/G  L  0  0  1  26  10  95  2  134  0.801  5.8-7.1  N  Ma  O/C/F/M/G  A  0  0  3  13  9  20  4  49  0.293  6.2 - 9.3  N  Ma  O/C/F/M/G  A  0  0  0  7  5  6  9  27  0.161  7.3 - 9.0  H  Ma  O/C/F/M/G  A  1  0  0  0  0  0  0  1  0.006  5.1 -6.5  N  Ma  F/C  A  0  0  0  1  1  5  0  7  0.042  3.5 - 5.2  H  Ma  G/F/M/C  A  0.102  Ma  G/M/O/R  L/A A  Pre-harvest Control 'Amara familiaris 'Amara farcta  (Duftschmid)  LeConte  Amara idahoana  (Casey)  Amara laevipennis 'Amara littoralis  Kirby Mannerheim  'Amara lunicollis 'Badister  Schiodte  obtusus  Trichocellus  LeConte"  cognatus  (Gyllenhal)  20%I.T.S. 20%P.C.  % of Total Length(mm)  'Harpalus  animosus  Casey  0  1  0  3  8  4  1  17  12.5-14.0  N  'Harpalus  fulvilabris  Mannerheim  0  0  1  0  0  0  0  1  0.006  8.8- 10.8  N  D  O/F  Harpalus nigritarsus  C R . Sahlberg  1  3  6  5  2  4  9  30  0.179  7.0 - 9.2  H  Ma  O/G/M  A  'Harpalus  innocuus  LeConte  0  0  2  0  1  2  1  6  0.036  8.2 - 9.7  N  Ma  O/M  A  'Harpalus  plenalis  0  0  0  0  0  0  1  1  0.006  7.5 - 8.4  N  Ma  O  L  0  0  1  4  3  5  6  19  0.114  7.5 - 9.3  N  Ma  O/F/G/M/C  L/A  0  0  0  0  0  0  0  0  0.000  8.8-10.4  H  Ma  O/F/M  L/A  0  0  0  0  2  0  0  2  0.012  7.3 - 9.8  H  Ma  O/C/G/M  L/A  68  492  511  356  233  374  386  2420  14.462  7.0-11.1  N  D  O/F/T/G/C/M  L/A  61  131  225  51  7  123  114  712  4.255  8.0-11.8  N  Ma  F/M/R  L  15  172  61  145  27  88  63  571  3.412  8.7-11.2  N  D  C/F  L  0  0  0  0  0  0  0  0  0.000  4.2 - 6.0  H  Ma  P/F  A  Casey  Harpalus opaclpennis Harpalus solitaris 'Harpalus  (Haldeman)  Dejean  fuscipalpis  Calathus ingratus Calathus advena  Sturm  Dejean (LeConte)  Synuchus  impunctatus  Sericoda  quadripunctata  (Say) (DeGeer)  Agonum retractum  LeConte  8  0  3  0  0  1  0  12  0.072  6.2 - 7.6  N  D  F  A  'Agonum  cupreum  Dejean  0  0  0  0  0  1  0  1  0.006  7.0-9.5  N  D  O/G/M/C  A  Cymindis cribicollis  Dejean  3  18  28  6  21  6  16  98  0.586  8.4-11.0  N  D  F/M/C  A  4  46  55  20  13  49  35  222  1.327  8.0 - 9.5  N  B  O/T/M  A  7  6  20  55  18  65  32  203  1.213  2.7-3.5  N  D  O/F/G/M/C  A  TOTAL  943  2903  3207  2484  1571  3016  2610  16734  % TOTAL  5.63  17.35  19.16  14.84  9.39  18.02  15.60  # SPECIES  22  19  27  30  31  30  30  49  # GENERA  14  12  14  15  13  14  14  22  # IMMIGRANT SPECIES (AFTER LOGGING)  10  12  13  12  11  28  # EMIGRANT SPECIES (AFTER LOGGING)  2  1  1  1  0  0  Cymindis unicolor Syntomus  Kirby  americanus  (Dejean)  Note: N=nearctic, H=holarctic, ^introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, 0=open, R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, A=adult, L=larval, P=pyrophilous, W.morph=wing morphology, 0/W=overwintering stage, Dist.= distribution, I.T.S.=individual tree selection, P.C=patch-cut, (R)=reserves, *=unique to Mud Lake, **=unique to pre-harvest.  Table  3: Total number of specimens caught in all experimental treatments at the Opax Mountain site (List of species as in Table 1). Pre-harvest Control  Trachypachus  holmbergi  'Leistus ferruginosus Notiophilus  Mannerheim  Mannerheim  Total 8  Dist.  W.morph.  Habitat  O/W Stage  0.031  3.8 -5.8  N  Ma  O  A  N  Ma  R/F  L  % of Total Length(mm)  2  0.008  6  108  148  125  81  165  82  715  2.731  4.8 -5.7  N  D  O/F/R  A  0  0  0  0  0  0  0  0  0.000  7.0 -8.5  H  Ma  R/C  A  Casey  0  0  0  0  0  0  0  0  0.000  14.0 -16.0  N  Ma  G  L  (LeConte)  0  0  0  0  0  0  0  0  0.000  15.0 -20.0  N  B  O/M  L  0  0  0  0  0  0  0  0  0.000  16.0 -24.0  N  D  O/F/M/C  A  49  1780  1591  1217  1254  1429  1342  8662  33.080  16.0 -23.0  N  B  O/F/M/G/C  A  relictus  (Fabricius)  LeConte  (G.H. Horn)  Scaphinotus  marginatus  (Fischer von Waldheim) (Fischer von Waldheim)  (Paykull) Erichson  "Trechus tenuiscapus  erasum  Bembidion  iridescens  Bembidion  grapii  Bembidion  rupicola  1  0  1  0  0  0  1  3  0.011  16.0 -20.0  N  B  F  L/A  0  86  8  17  63  12  20  206  0.787  17.0 -24.0  N  B  F/R/C  L/A  18  124  153  70  97  83  191  736  2.811  11.5 -19.0  N  B  F/R/C/T  L  3  7  5  8  1  8  0  32  0.122  6.2 -9.8  H  Ma  O/F/M  A  3.6 -4.1  I  D  O/M/C/F  L/A  0  0  0  0  0  0  0  0  0.000  Lindroth"  2  0  0  0  0  0  0  2  0.008  4.1 -4.9  N  B  F  A  LeConte  76  17  22  33  9  98  28  283  1.081  2.9 -3.8  N  D  O/F  A  0  0  0  0  0  1  0  1  0.004  3.0 -4.2  N  Ma  R/F  A  0  0  0  0  0  0  0  0  0.000  3.8 -4.8  N  Ma  R/F  A  13  1  32  1  54  0.206  4.0 -4.8  H  Po  O/M/C/F  A  dyschirinum  'Bembidion  'Patrobus  1 0  angusticollis  Bembidion  2 0  Say  Trechus obtusus  0 1  Casey  arctica  50%I.T.S. 50%P.C. 50%I.T.S.(R)  0  Scaphinotus  Miscodera  5  0  Carabus taedatus agassii 'Scaphinotus  0  1  Cicindela nebraskana  Carabus serratus  20%I.T.S. 20%P.C.  0  direclus  wilkesii  0  7.8 -9.3  Loricera pilicornis pilicornis  Calosoma  0  LeConte (LeConte)  Gyllenhal  5  1  1  (Kirby)  0  0  0  0  0  1  0  1  0.004  4.1 -5.5  N  Ma  O/F/C/R/M  A  Chaudoir  0  0  0  0  0  0  1  1  0.004  8.3- 12.0  N  Ma  R/O  L/A  stygicus  Stereocerus  haematopus  (Dejean)  0  0  0  0  0  0  0  0  0.000  9.5- 12.5  H  D  T  A  Pterostichus  adstrictus  Eschscholtz  47  67  97  205  335  273  183  1207  4.610  9.5- 13.0  H  Ma  G/O/F/C/R  L/A  Pterostichus  oregonus  LeConte  2  1  0  1  0  0  28  32  0.122  12.0 -15.0  N  Ma  F  A  Pterostichus  pensylvanicus  3  1  19  2  2  1  6  34  0.130  9.5- 12.0  N  D  F/R  A  Pterostichus  melanarius  0  0  0  0  0  0  0  0  0.000  12.0 -19.0  I  D  C/F/M/R/G  L  Pterostichus  neobrunneus  153  1086  752  1353  1543  617  813  6317  24.124  11.5 -14.2  N  B  F  A  0  0  0  0  4  0  0  4  0.015  6.5 -8.0  N  B  R/F  A L  LeConte (llliger) Lindroth  'Pterostichus  riparius  (Dejean)  Amara iatior  (Kirby)  0  0  0  0  0  2  0  2  0.008  7.8- 10.8  N  Ma  O/F/C/M  Amara obesa  (Say)  0  0  0  0  0  0  0  0  0.000  9.0- 12.5  N  D  O/R/C/M  L  0.004  6.2 -8.8  I  Ma  M/O/G/C  A  N  Ma  ?  ?  'Amara aenea  (DeGeer)  0  0  0  0  0  0  1  1  Amara conflata  LeConte  0  0  0  0  0  0  0  0  0.000  7.6- 10.0  0  0  0  3  0  16  3  22  0.084  7.0 -9.0  N  Ma  O/M/G/C  A  0  0  0  0  0  0  0  0  0.000  6.5 -8.7  H  Ma  O/G/M/F  A  Amara ellipsis Amara erratica  (Casey) (Duftschmid)  Table 3: Total number of specimens caught in all experimental treatments at the Opax Mountain site (cont.). Dist.  W.morph.  Habitat  O/W Stage  0  0  0  0  0  0  0  0  0.000  5.6  7.2  I  Ma  O/C/F/M  A  0  0  0  0  0  0  0  0  0.000  6.5  9.0  N  Ma  O/R/C/M  A  1  2  3  3  6  45  5  65  0.248  5.0  6.5  N  D  O/M/G  L  Kirby  0  1  0  9  0  27  2  39  0.149  5.8  7.1  N  Ma  O/C/F/M/G  A  Mannerheim  0  0  0  0  0  0  0  0  0.000  6.2  9.3  N  Ma  O/C/F/M/G  A  0  0  0  0  0  0  0  0  0.000  7.3  9.0  H  Ma  O/C/F/M/G  A  0  0  0  0  0  0  0  0  0.000  5.1  6.5  N  Ma  F/C  A  1  0  0  0  0  0  0  1  0.004  3.5  5.2  H  Ma  G/F/M/C  A  14.0  N  Ma  G/M/O/R  L/A  Pre-harvest Control Amara familiaris Amara tarda  (Duftschmid)  LeConte  Amara idahoana  (Casey)  Amara laevipennis Amara littoralis Amara lunicollis  Schiodte  Badister obtusus Trichocellus  LeConte  cognatus  (Gyllenhal)**  20%I.T.S. 20%P.C.  50%I.T.S. 50%P.C. 50%I.T.S.(R)  Total  % of Total Length(mm)  Harpalus animosus  Casey  0  0  0  0  0  0  0  0  0.000  12.5  Harpalus  Mannerheim  0  0  0  0  0  0  0  0  0.000  8.8- 10.8  N  D  O/F  A  fulvilabris  Harpalus nigritarsus  C R . Sahlberg  Harpalus innocuus Harpalus plenalis Harpalus  LeConte Casey  opacipennis  "Harpalus  solilaris  Harpalus  luscipalpis  Calathus ingratus Calathus advena  (Haldeman) Dejean Sturm  Dejean (LeConte)  Synuchus  impunctatus  "Sericoda  quadripunctata  Agonum  retractum  Agonum cupreum Cymindis  cribicollis  Cymindis  unicolor  Syntomus  (Say) (DeGeer)  LeConte Dejean Dejean Kirby  americanus  (Dejean)  1  4  21  5  12  24  6  73  0.279  7.0  9.2  H  Ma  O/G/M  A  0  0  0  0  0  0  0  0  0.000  8.2  9.7  N  Ma  O/M  A  0  0  0  0  0  0  0  0  0.000  7.5  8.4  N  Ma  0  L  0  0  0  0  0  7  0  7  0.027  7.5  9.3  N  Ma  O/F/G/M/C  L/A  0  1  0  0  0  0  0  1  0.004  8.8- 10.4  H  Ma  O/F/M  L/A  7.3  0  0  0  0  0  0  0  0  0.000  9.8  H  Ma  O/C/G/M  L/A  19  227  663  408  508  478  271  2574  9.830  7.0- 11.1  N  D  O/F/T/G/C/M  L/A  122  796  458  690  927  328  1135  4456  17.017  8.0- 11.8  N  Ma  F/M/R  L  0  10  39  9  16  29  47  150  0.573  8.7- 11.2  N  D  C/F  L  0  0  0  1  0  0  0  1  0.004  4.2  6.0  H  Ma  P/F  A  0  0  0  1  1  0  1  3  0.011  6.2  7.6  N  D  F  A  0  0  0  0  0  0  0  0  0.000  7.0  9.5  N  D  O/G/M/C  A  2  7  14  6  3  2  16  50  0.191  8.4- 11.0  N  D  F/M/C  A  5  42  60  90  65  69  50  381  1.455  8.0  9.5  N  B  O/T/M  A  0  2  8  11  2  28  8  59  0.225  2.7  3.5  N  D  O/F/G/M/C  A  26185  TOTAL  516  4371  4063  4285  4931  3777  4242  % TOTAL  1.97  16.69  15.52  16.36  18.83  14.42  16.20  # SPECIES  19  22  19  24  21  25  25  36  #GENERA  12  13  12  15  14  13  14  19  # IMMIGRANT S P E C I E S (AFTER LOGGING)  1  4  2  6  6  14  # EMIGRANT SPECIES (AFTER LOGGING)  5  2  3  3  3  1  Note: N=nearotic, H=holarctic, ^introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, O=open; R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, L= larval, A=adult, P=pyrophilous, W.morph=wing morphology, 0/W=overwintering stage, Dist.= distribution, I.T.S.=individual tree selection, P.C.=patch-cut, (R)=reserves, *=unique to Opax Mtn, **=unique to pre-harvest.  Table  4: Total number of specimens caught in all experimental treatments (spring samples) at Opax Mountain Silvicultural Systems Research Project (List of species as in Table 1) .  Trachypachus  holmbergi  Mannerheim  Leistus ferruginosus  Mannerheim  Notiophilus  Casey  directus  Loricera pilicornis pilicornis Cicindela  Carabus  Total  % of Total  Length(mm)  Habitat  O/W Stage  13  31  0.138  3.8 -5.8  N  Ma  O  A  1  0  0  0  0  0  1  0.004  7.8 -9.3  N  Ma  R/F  L  41  50  56  36  90  29  302  1.344  4.8 -5.7  N  D  O/F/R  A  Distribution W.morph.  0  0  0  0  0  0.000  7.0 -8.5  H  Ma  R/C  A  0  1  0  0  1  0.004  14.0 -16.0  N  Ma  G  L  (LeConte)  0  0  0  0  0  1  1  0.004  15.0 -20.0  N  B  O/M  L  Say  0  0  0  1  0  20  21  0.093  16.0 -24.0  N  D  O/F/M/C  A  1003  915  666  638  746  653  4621  20.562  16.0 -23.0  N  B  O/F/M/G/C  A  Scaphinotus  angusticollis  Scaphinotus  marginatus arctica  LeConte  (G.H. Horn) (Fischer von Waldheim) (Fischer von Waldheim)  (Paykull) Erichson  Trechus tenuiscapus  Lindroth  Bembidion  dyschirinum  Bembidion  erasum  Bembidion  iridescens  Bembidion  grapii  Bembidion  rupicola  LeConte  LeConte  0  0  0  0  0  0  0  0.000  16.0 -20.0  N  B  F  L/A  18  10  2  5  2  5  42  0.187  17.0 -24.0  N  B  F/R/C  L/A  55'  72  43  35  55  86  346  1.540  11.5 -19.0  N  B  F/R/C/T  L  5  4  3  2  8  0  22  0.098  6.2 -9.8  H  Ma  O/F/M  A  0  0  0  0  0  0  0  0.000  3.6 -4.1  I  D  O/M/C/F  L/A  0  0  0  0  0  0  0  0.000  4.1 -4.9  N  B  F  A  16  26  62  19  126  34  283  1.259  2:9 -3.8  N  D  O/F  A  3.0 -4.2  N  Ma  R/F  A  Ma  R/F  A A  0  0  0  0  1  0  1  0.004  0  0  3  0  0  0  3  0.013  3.8 -4.8  N  Gyllenhal  0  1  13  1  25  0  40  0.178  4.0 -4.8  H  Po  O/M/C/F  (Kirby)  0  1  0  0  1  0  2  0.009  4.1 -5.5  N  Ma  O/F/C/R/M  A  Chaudoir  0  0  0  0  0  1  1  0.004  8.3- 12.0  N  Ma  R/O  L/A  stygicus  (LeConte)  Stereocerus  haematopus  Pterostichus  adstrictus  Eschscholtz  Pterostichus  oregonus  LeConte  Pterostichus  pensylvanicus  Pterostichus  melanarius  Pterostichus  neobrunneus  Pterostichus  riparius  Amara latior  50%I.T.S.(R)  10  0  relictus  Patrobus  50%P.C.  0  0  taedatus agassii  Trechus obtusus  50%I.T.S.  6  0  Scaphinotus  Miscodera  20%P.C.  1  0  wilkesii  Carabus serratus  20%I.T.S.  1  Casey  nebraskana  Calosoma  (Fabricius)  Control  (Dejean)  LeConte (llliger) Lindroth  (Dejean)  0  0  0  0  0  1  0.004  9.5- 12.5  H  D  T  A  266  307  397  426  648  627  2671  11.885  9.5- 13.0  H  Ma  G/O/F/C/R  L/A  2  5  66  20  2  64  159  0.707  12.0 -15.0  N  Ma  F  A  2  14  2  2  1  2  23  0.102  9.5- 12.0  N  D  F/R  A  0  0  0  0  0  0  0  0.000  12.0 -19.0  I  D  C/F/M/R/G  L  1326  1308  1434  1485  926  937  7416  32.998  11.5 -14.2  N  B  F  A  0  0  0  3  0  0  3  0.013  6.5 -8.0  N  B  R/F  A  Ma  O/F/C/M  L  1  (Kirby)  0  0  0  0  1  0  1  0.004  7.8- 10.8  N  Amara obesa  (Say)  0  0  0  0  0  0  0  0.000  9.0- 12.5  N  D  O/R/C/M  L  Amara aenea  (DeGeer)  0  0  0  0  0  1  1  0.004  6.2 -8.8  I  Ma  M/O/G/C  A  0  0  1  0  0  0  1  0.004  7.6- 10.0  N  Ma  ?  ?  0  0  8  7  38  1  54  0.240  7.0 -9.0  N  Ma  O/M/G/C  A  0  0  1  1  1  0  3  0.013  6.5 -8.7  H  Ma  O/G/M/F  A  Amara conflata Amara ellipsis Amara erratica  LeConte (Casey) (Duftschmid)  Table  4: Total number of specimens caught in all experimental treatments (spring samples) at Opax Mountain Silvicultural Systems Research Project (cont.). Control  20%I.T.S.  20%P.C.  50%I.T.S.(R)  Total  Habitat  O/W Stage  Amara familiaris (Duftschmid)  0  0  0  0  1  1  2  0.009  5.6 - 7.2  I  Ma  O/C/F/M  A  Amara farcta LeConte  0  1  0  0  0  0  1  0.004  6.5-9.0  N  Ma  O/R/C/M  A  Amara idahoana (Casey)  0  0  2  1  1  0  4  0.018  5.0-6.5  N  D  O/M/G  L  Amara laevipennis Kirby  1  2  31  8  105  4  151  0.672  5.8-7.1  N  Ma  O/C/F/M/G  A  Amara littoralis Mannerheim  0  1  10  7  18  3  39  0.174  6.2 - 9.3  N  Ma  O/C/F/M/G  A  Amara lunicollis Schiodte  0  0  7  5  6  7  25  0.111  7.3 - 9.0  H  Ma  O/C/F/M/G  A  Badister obtusus LeConte  0  0  0  0  0  0  0  0.000  5.1 -6.5  N  Ma  F/C  A  Trichocellus cognatus (Gyllenhal)  0  0  0  0  5  0  5  0.022  3.5-5.2  H  Ma  G/F/M/C  A  Harpalus animosus Casey  1  0  2  7  3  1  14  0.062  12.5-14.0  N  Ma  G/M/O/R  L/A  Harpalus fulvilabris Mannerheim  0  0  0  0  0  0  0  0.000  8.8-10.8  N  D  O/F  A  Harpalus nigritarsus C R . Sahlberg  7  16  8  10  21  13  75  0.334  7.0 - 9.2  H  Ma  O/G/M  A  Harpalus innocuus LeConte  0  2  0  1  2  1  6  0.027  8.2 - 9.7  N  Ma  O/M  A  Harpalus plenalis Casey  0  0  0  0  0  1  1  0.004  7.5 - 8.4  N  Ma  O  L  Harpalus opacipennis (Haldeman)  0  1  2  3  7  4  17  0.076  7.5 - 9.3  N  Ma  O/F/G/M/C  L/A  Harpalus solitaris Dejean  1  0  0  0  0  0  1  0.004  8.8-10.4  H  Ma  O/F/M  L/A  Harpalus fuscipalpis Sturm  0  0  0  1  0  0  1  0.004  7.3 - 9.8  H  Ma  O/C/G/M  L/A  Calathus ingratus Dejean  314  574  409  377  442  246  2362  10.510  7.0-11.1  N  D  O/F/T/G/C/M  L/A  Calathus advena (LeConte)  638  472  486  612  281  797  3286  14.621  8.0-11.8  N  Ma  F/M/R  L  Synuchus impunctatus (Say)  0  0  0  0  0  0  0  0.000  8.7-11.2  N  D  C/F  L  Sericoda quadrlpunctata (DeGeer)  0  0  0  0  0  0  0  0.000  4.2 - 6.0  H  Ma  P/F  A  Agonum retractum LeConte  0  3  1  1  1  0  6  0.027  6.2 - 7.6  N  D  F  A  Agonum cupreum Dejean  0  0  0  0  1  0  1  0.004  7.0 - 9.5  N  D  O/G/M/C  A  F/M/C  A  Cymindis cribicollis Dejean Cymindis unicolor Kirby Syntomus americanus (Dejean)  .  50%I.T.S. 50%P.C.  % of Total Length(mm)  Distribution W.morph.  4  1  3  1  6  17  0.076  8.4-11.0  N  D  30  31  37  33  40  39  210  0.934  8.0 - 9.5  N  B  O/T/M  A  4  20  62  11  69  33  199  0.885  2.7-3.5  N  D  O/F/G/M/C  A  22474  2  TOTAL  3734  3841  3821  3762  3685  3631  % TOTAL  16.61  17.09  17.00  16.74  16.40  16.16  # SPECIES  21  25  28  31  34  30  #GENERA  13  13  13  13  14  14  19  # IMMIGRANT SPECIES (AFTER LOGGING)  7  9  13  15  12  27  # EMIGRANT SPECIES (AFTER LOGGING)  3  2  3  2  3  1  48  Note: N=nearctic, H=holarctic, l=introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, O=open, R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, A=adult, L=larval. P=pyrophilous, W.morph=wing morphology, O/W=0verwintering stage, I.T.S.=individual tree selection, P.C.=patch-cut, (R)=reserves.  Table 5 : Total number of specimens caught in all experimental treatments (spring samples) at Mud Lake site (List of species as in Table 1).  Trachypachus  holmbergi  Mannerheim  Control  20%I.T.S.  20%P.C.  Total  % of Total  Length(mm)  Distribution  W.morph.  Habitat  O/W Stage  1  1  2  0  8  12  24  0.320  3.8 - 5.8  N  Ma  O  A  50%I.T.S. 50%P.C. 50%I.T.S.(R)  Leistus ferruginosus  Mannerheim  0  0  0  0  0  0  0  0.000  7.8 - 9.3  N  Ma  R/F  L  Notiophilus  Casey  3  12  6  4  10  5  40  0.533  4.8 - 5.7  N  D  O/F/R  A  directus  Loricera pilicornis pilicornis 'Cicindela 'Calosoma 'Carabus  0  0  0  0  0  0  0  0.000  7.0-8.5  H  Ma  R/C  A  0  0  0  1  0  0  1  0.013  14.0-16.0  N  Ma  G  L  (LeConte)  0  0  0  0  0  1  1  0.013  15.0-20.0  N  B  O/M  L  0  0  0  1  0  20  21  0.280  16.0-24.0  N  D  O/F/M/C  A  276  249  170  165  148  146  1154  15.379  16.0-23.0  N  B  O/F/M/G/C  A  0  0  0  0  0  0  0  0.000  16.0^20.0  N  B  F  L/A  2  9  0  0  0  0  11  0.147  17.0-24.0  N  B  F/R/C  L/A  22  33  10  3  28  16  112  1.493  11.5-19.0  N  B  F/R/C/T  L  (Paykull)  0  1  0  0  0  0  1  0.013  6.2 - 9.8  H  Ma  O/F/M  A  Erichson  0  0  0  0  0  0  0  0.000  3.6-4.1  I  D  O/M/C/F  L/A  wilkesii serratus  Say  Carabus taedatus  agassii  Scaphinotus  relictus  Scaphinotus  angusticollis  Scaphinotus  marginatus  Miscodera  arctica  Trechus obtusus  Trechus tenuiscapus  0  0  0  0  0  0  0.000  4.1 -4.9  N  B  F  A  5  10  40  12  48  14  129  1.719  2.9-3.8  N  D  O/F  A  0  0  0  0  0  0  0  0.000  3.0 - 4.2  N  Ma  R/F  A  0  0  3  0  0  0  3  0.040  3.8 - 4.8  N  Ma  R/F  A  Gyllenhal  0  0  1  0  9  0  10  0.133  4.0 - 4.8  H  Po  O/M/C/F  A  (Kirby)  0  1  0  0  0  0  1  0.013  4.1 -5.5  N  Ma  O/F/C/R/M  A  0  0  0  0  0  0  0  0.000  8.3-12.0  N  Ma  R/O  L/A  0  0  0  0  0  1  1  0.013  9.5-12.5  H  D  T  A  213  222  238  148  449  494  1764  23.508  9.5-13.0  H  Ma  G/O/F/C/R  L/A  2  5  65  20  2  47  141  1.879  12.0-15.0  N  Ma  F  A  1  0  0  0  0  2  3  0.040  9.5-12.0  N  D  F/R  A  0  0  0  0  0  0  0  0.000  12.0-19.0  I  D  C/F/M/R/G  L  erasum  LeConte  LeConte  iridescens  Bembidion  rupicola  Patrobus stygicus 'Stereocerus Pterostichus  (Fischer von Waldheim)  0  Bembidion  grapii  (Fischer von Waldheim)  Lindroth  dyschirinum  Bembidion  LeConte  (G.H. Horn)  Bembidion  'Bembidion  (Fabricius)  Casey  nebraskana  (LeConte)  Chaudoir  haematopus  (Dejean)  adstrictus  Eschscholtz  Pterostichus  oregonus  LeConte  Pterostichus  pensylvanicus  Pterostichus  melanarius  Pterostichus  neobrunneus  Pterostichus  riparius  LeConte (llliger) Lindroth  428  679  326  196  412  318  2359  31.437  11.5-14.2  N  B  F  A  0  0  0  0  0  0  0  0.000  6.5 - 8.0  N  B  R/F  A  Amara latior (Kirby)  0  0  0  0  0  0  0  0.000  7.8-10.8  N  Ma  O/F/C/M  L  Amara obesa  (Say)  0  0  0  0  0  0  0  0.000  9.0-12.5  N  D  O/R/C/M  L  Amara aenea  (DeGeer)  0  0  0  0  0  0  0  0.000  6.2 - 8.8  I  Ma  M/O/G/C  A  0  0  1  0  0  0  1  0.013  7.6-10.0  N  Ma  0  0  6  7  25  0  38  0.506  7.0 - 9.0  N  Ma  O/M/G/C  A  0  0  0  1  1  0  2  0.027  6.5 - 8.7  H  Ma  O/G/M/F  A  'Amara conflata Amara ellipsis 'Amara erratica  (Dejean)  LeConte (Casey) (Duftschmid)  ?  ?  Table 5: Total number of specimens caught in all experimental treatments (spring samples) at Mud Lake site (cont.). Control  20%I.T.S.  20%P.C.  Total  % of Total  Length(mm)  Distribution  W.morph.  Habitat  O/W Stage  "Amara familiaris (Duftschmid)  0  0  0  0  1  1  2  0.027  5.6 - 7.2  I  Ma  O/C/F/M  A  'Amara tarda LeConte  0  1  0  0  0  0  1  0.013  6.5 - 9.0  N  Ma  O/R/C/M  A  Amara idahoana  0  0  0  0  0  0  0  0.000  5.0-6.5  N  D  O/M/G  L  Amara laevipennis Kirby  0  2  24  8  82  2  118  1.570  5.8-7.1  N  Ma  O/C/F/M/G  A  'Amara littoralis Mannerheim  0  1  10  7  18  3  39  0.520  6.2 - 9.3  N  Ma  O/C/F/M/G  A  'Amara lunicollis Schiodte  0  0  7  5  6  7  25  0.333  7.3 - 9.0  H  Ma  O/C/F/M/G  A  'Badister obtusus LeConte  0  0  0  0  0  0  0  0.000  5.1 -6.5  N  Ma  F/C  A  Trichocellus cognatus (Gyllenhal)  0  0  0  0  5  0  5  0.067  3.5-5.2  H  Ma  G/F/M/C  A  'Harpalus animosus Casey  1  0  2  7  3  1  14  0.187  12.5-14.0  N  Ma  G/M/O/R  L/A  Harpalus fulvilabris Mannerheim  0  0  0  0  0  0  0  0.000  8.8-10.8  N  D  O/F  A  Harpalus nigritarsus C R . Sahlberg  3  4  4  1  4  9  25  0.333  7.0 - 9.2  H  Ma  O/G/M  A  'Harpalus innocuus LeConte  0  2  0  1  2  1  6  0.080  8.2 - 9.7  N  Ma  O/M  A  'Harpalus plenalis Casey  0  0  0  0  0  1  1  0.013  7.5 - 8.4  N  Ma  O  L  Harpalus opadpennis  0  1  2  3  2  4  12  0.160  7.5 - 9.3  N  Ma  O/F/G/M/C  L/A  Harpalus solitaris Dejean  0  0  0  0  0  0  0  0.000  8.8-10.4  H  Ma  O/F/M  L/A  'Harpalus fusdpalpis Sturm  0  0  0  1  0  0  1  0.013  7.3 - 9.8  H  Ma  O/C/G/M  L/A  201  194  145  76  155  92  863  11.501  7.0-11.1  N  D  O/F/T/G/C/M  L/A  69  132  22  2  57  46  328  4.371  8.0-11.8  N  Ma  F/M/R  L  Synuchus impundatus (Say)  0  0  0  0  0  0  0  0.000  8.7-11.2  N  D  C/F  L  Sericoda quadripundata  0  0  0  0  0  0  0  0.000  4.2 - 6.0  H  Ma  P/F  A  Agonum retractum LeConte  0  3  0  0  1  0  4  0.053  6.2 - 7.6  N  D  F  A  'Agonum cupreum Dejean  0  0  0  0  1  0  1  0.013  7.0 - 9.5  N  D  O/G/M/C  A  8.4-11.0  N  D  F/M/C  A  (Casey)  (Haldeman)  Calathus ingratus Dejean Calathus advena  (LeConte)  (DeGeer)  Cymindis cribicollis Dejean  2  3  Cymindis unicolor Kirby  15  13  Syntomus americanus (Dejean)  3  14  50%I.T.S. 50%P.C. 50%I.T.S.(R)  3  1  10  6  20  15  79  1.053  8.0 - 9.5  N  B  O/T/M  A  52  10  44  28  151  2.012  2.7-3.5  N  D  O/F/G/M/C  A  7504  0  3  12  TOTAL  1247  1592  1146  688  1542  1289  % TOTAL  16.62  21.22  15.27  9.17  20.55  17.18  # SPECIES  17  23  22  24  27  26  #GENERA  10  13  11  11  13  13  17  # IMMIGRANT SPECIES (AFTER LOGGING)  8  8  10  12  10  23  # EMIGRANT SPECIES (AFTER LOGGING)  2  3  3  2  1  0  0.160  40  Note: N=nearctic, H=holarctic, l=introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, 0=open, R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, A=adult, L=larval, P=pyrophilous, W.morph=wing morphology, 0/W=overwintering stage, I.T.S.=individual tree selection, P.C.=patch-cut, (R)=reserves, *=unique to Mud Lake.  Table  6: Total number of specimens caught in all experimental treatments (spring samples) at Opax Mtn. site (List of species as in Table 1). 50%I.T.S.(R)  Total  % of Total  Length(mm)  Distribution  W.morph.  Habitat  O/W Stage  Trachypachus holmbergi Mannerheim  0  0  4  0  2  1  7  0.047  3.8 - 5.8  N  Ma  O  A  'Leistus ferruginosus Mannerheim  1  0  0  0  0  0  1  0.007  7.8 - 9.3  N  Ma  R/F  L  38  38  50  32  80  24  262  1.750  4.8 - 5.7  N  D  O/F/R  A  Loricera pilicornis pilicornis (Fabricius)  0  0  0  0  0  0  0  0.000  7.0 - 8.5  H  Ma  R/C  A  Cicindela nebraskana  Casey  0  0  0  0  0  0  0  0.000  14.0-16.0  N  Ma  G  L  Calosoma wilkesii (LeConte)  0  0  0  0  0  0  0  0.000  15.0-20.0  N  B  O/M  L  Carabus serratus Say  0  0  0  0  0  0  0  0.000  16.0-24.0  N  D  O/F/M/C  A  Control  Notiophilus directus Casey  Carabus taedatus agassii LeConte  20%I.T.S. 20%P.C.  50%I.T.S. 50%P.C.  727  666  496  473  598  507  3467  23.160  16.0-23.0  N  B  O/F/M/G/C  A  Scaphinotus relictus (G.H. Horn)  0  0  0  0  0  0  0  0.000  16.0-20.0  N  B  F  L/A  Scaphinotus angusticollis (Fischer von Waldheim)  16  1  2  5  2  5  31  0.207  17.0-24.0  N  B  F/R/C  L/A  Scaphinotus marginatus (Fischer von Waldheim)  33  39  33  32  27  70  234  1.563  11.5-19.0  N  B  F/R/C/T  L  5  3  3  2  8  0  21  0.140  6.2 - 9.8  H  Ma  O/F/M  A  0  0  0  0  0  0  0  0.000  3.6-4.1  I  D  O/M/C/F  L/A  Miscodera arctica  (Paykull)  Trechus obtusus Erichson Trechus tenuiscapus Lindroth  0  0  0  0  0  0  0  0.000  4.1 -4.9  N  B  F  A  Bembidion dyschihnum LeConte  11  16  22  7  78  20  154  1.029  2.9-3.8  N  D  O/F  A  'Bembidion erasum LeConte  0  0  0  0  1  0  1  0.007  3.0 - 4.2  N  Ma  R/F  A  Ma  R/F  A A  Bembidion iridescens (LeConte)  0  Bembidion grapii Gyllenhal Bembidion rupicola (Kirby) 'Patrobus stygicus Chaudoir  0  0  0  0  0  0  0.000  3.8 - 4.8  N  0  1  0  0  12  1  16  0  30  0.200  4.0 - 4.8  H  Po  O/M/C/F  0  0  1  0  1  0.007  4.1 -5.5  N  Ma  O/F/C/R/M  0  0  A  0  0  0  1  1  0.007  8.3-12.0  N  Ma  R/O  L/A  Stereocerus haematopus (Dejean)  0  0  0  0  0  0  0  0.000  9.5-12.5  H  D  T  A  Pterostichus adstrictus Eschscholtz  53  85  159  278  199  133  907  6.059  9.5-13.0  H  Ma  G/O/F/C/R  L/A  Pterostichus oregonus LeConte  0  0  1  0  0  17  18  0.120  12.0-15.0  N  Ma  F  A  Pterostichus pensylvanicus LeConte  1  14  2  2  1  0  20  0.134  9.5-12.0  N  D  F/R  A  Pterostichus melanarius (llliger)  0  0  0  0  0  0  0  0.000  12.0-19.0  I  D  C/F/M/R/G  L  898  629  1108  1289  514  619  5057  33.781  11.5-14.2  N  B  F  A  0  0  0  3  0  0  3  0.020  6.5 - 8.0  N  B  R/F  A  Ma  O/F/C/M  L  Pterostichus neobrunneus Lindroth 'Pterostichus riparius (Dejean) Amara latior (Kirby)  0  0  0  0  1  0  1  0.007  7.8-10.8  N  Amara obesa (Say)  0  0  0  0  0  0  0  0.000  9.0-12.5  N  D  O/R/C/M  L  'Amara aenea  (DeGeer)  0  0  0  0  0  1  1  0.007  6.2 - 8.8  I  Ma  M/O/G/C  A  Amara conflata LeConte  0  0  0  0  0  0  0  0.000  7.6-10.0  N  Ma  ?  ?  Amara ellipsis (Casey)  0  0  2  0  13  1  16  0.107  7.0 - 9.0  N  Ma  O/M/G/C  A  Amara erratica (Duftschmid)  0  0  0  0  0  0  0  0.000  6.5 - 8.7  H  Ma  O/G/M/F  A  Table 6: Total number of specimens caught in all experimental treatments (spring samples) at Opax Mtn. Site (cont.). Control Amara familiaris Amara farcla  (Duftschmid)  50%I.T.S.(R)  Total  % of Total  Length(mm)  Distribution W.morph.  Habitat  O/W Stage  0  0  0  0  0  0  0.000  5.6 -7.2  I  Ma  O/C/F/M  A  0  0  0  0  0  0  0  0.000  6.5 -9.0  N  Ma  O/R/C/M  A  0  0  2  1  1  0  4  0.027  5.0 -6.5  N  D  O/M/G  L  Kirby  1  0  8  0  23  2  34  0.227  5.8 -7.1  N  Ma  O/C/F/M/G  A  Mannerheim  0  0  0  0  0  0  0  0.000  6.2 -9.3  N  Ma  O/C/F/M/G  A  0  0  0  0  0  0  0  0.000  7.3 -9.0  H  Ma  O/C/F/M/G  A  0  0  0  0  0  0  0  0.000  5.1 -6.5  N  Ma  F/C  A  0  0  0  0  0  0  0  0.000  3.5 -5.2  H  Ma  G/F/M/C  A  0  0.000  12.5 -14.0  N  Ma  G/M/O/R  L/A  Amara idahoana  (Casey)  Amara laevipennis  Amara lunicollis  Schiodte  Badister obtusus Trichocellus  50%I.T.S. 50%P.C  0  LeConte  Amara littoralis  20%I.T.S. 20%P.C.  LeConte  cognatus  (Gyllenhal)  Harpalus animosus  Casey  0  0  0  0  0  0  Harpalus fulvilabris  Mannerheim  0  0  0  0  0  0  0  0.000  8.8- 10.8  N  D  O/F  A  4  12  4  9  17  4  50  0.334  7.0 -9.2  H  Ma  O/G/M  A  0  0  0  0  0  0  0  0.000  8.2 -9.7  N  Ma  O/M  A  0  0  0  0  0  0  0  0.000  7.5 -8.4  N  Ma  O  L  0  0  0  0  5  0  5  0.033  7.5 -9.3  N  Ma  O/F/G/M/C  L/A  Harpalus nigritarsus  C R . Sahlberg  Harpalus innocuus  LeConte  Harpalus plenalis  Casey  Harpalus opacipennis 'Harpalus  solitaris  (Haldeman)  Dejean  Harpalus fuscipalpis Calathus ingratus Calathus advena Synuchus  Sturm Dejean  (LeConte)  impunctatus  (Say)  Sericoda quadripunctata Agonum retractum Agonum cupreum  LeConte Dejean  Cymindis cribicollis Cymindis unicolor  (DeGeer)  Dejean Kirby  Syntomus americanus  (Dejean)  1  0  0  0  0  0  1  0.007  8.8- 10.4  H  Ma  O/F/M  L/A  0  0  0  0  0  0  0  0.000  7.3 - 9.8  H  Ma  O/C/G/M  L/A  113  380  264  301  287  154  1499  10.013  7.0- 11.1  N  D  O/F/T/G/C/M  L/A  569  340  464  610  224  751  2958  19.760  8.0- 11.8  N  Ma  F/M/R  L  0  0.000  8.7- 11.2  N  D  C/F  L  Ma  P/F  A  0  0  0  0  0  0  0  0  0  0  0  0  0  0.000  4.2 -6.0  H  0  0  1  1  0  0  2  0.013  6.2 -7.6  N  D  F  A  0  0  0  0  0  0  0  0.000  7.0 -9.5  N  D  O/G/M/C  A  0  1  1  0  0  3  5  0.033  8.4- 11.0  N  D  F/M/C  A  15  18  27  27  20  24  131  0.875  8.0 -9.5  N  B  O/T/M  A  1  6  10  1  25  5  48  0.321  2.7 -3.5  N  D  O/F/G/M/C  A  14970  TOTAL  2487  2249  2675  3074  2143  2342  % TOTAL  16.61  15.02  17.87  20.53  14.32  15.64  # SPECIES  17  16  22  18  23  19  31  #GENERA  12  10  13  12  12  12  15  # IMMIGRANT SPECIES (AFTER LOGGING)  2  7  4  8  6  14  # EMIGRANT SPECIES (AFTER LOGGING)  3  2  3  2  4  1  Note: N=nearctic, H=holarctic, ^introduced, Ma=macropterous, B=brachypterous, D=dimporphic, Po=polymorphic, 0=open, R=riparian, F=forest, G=grassland, M=meadow, C=cultivated, T=tundra, A=adult, L=larval, P=pyrophilous, W.morph=wing morphology, 0/W=overwintering stage, I.T.S.=individual tree selection, P.C.=patch-cut, (R)=reserves, *=unique to Opax Mtn.  Table 7: Abundance, species number, richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Mud Lake, based on total trap numbers for spring samples, 1995-1997.  N No. Species ES(100) H' 1/D'  Control 1247 17 8.27 1.695 4.446  20% I.T.S. 1592 23 10.00 1.752 4.036  20% P.C. 1146 22 13.07 2.097 5.917  50% I.T.S. 688 24 13.31 1.957 5.051  50% P.C. 1542 27 14.16 2.115 5.504  50% I.T.S.(R) 1289 26 13.14 1.918 4.367  Table 8: Abundance, species number, richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Mud Lake, based on total trap numbers for the entire season, 1993-1997.  N No. Species ES(100) H' 1/D'  Control 2903 19 10.22 1.946 5.398  20% I.T.S. 3207 27 11.67 2.035 5.734  20% P.C. 2484 30 14.79 2.247 6.573  50% I.T.S. 1571 31 14.48 2.027 4.802  50% P.C. 3016 30 14.90 2.252 6.776  50% I.T.S.(R) 2610 30 13.79 2.114 5.919  Table 9: Abundance, species number, richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Opax Mountain, based on total trap numbers for spring samples, 1995-1997.  N No. Species ES(100) H' 1/D'  Control 2487 17 8.27 1.534 3.691  20% I.T.S. 2249 16 9.11 1.734 4.562  20% P.C. 2675 22 9.53 1.712 4.005  50% I.T.S. 3074 18 7.87 1.623 3.896  50% P.C. 2143 23 12.19 2.040 5.685  50% I.T.S.(R) 2342 19 9.29 1.730 4.386  Table 10: Abundance, species number, richness, Shannon-Wiener and Simpson's diversity indices, for the six experimental treatments at Opax Mountain, based on total trap numbers for the entire season, 1993-1997.  N No. Species ES(100) H' 1/D'  Control 4371 22 9.29 1.635 3.768  20% I.T.S. 4063 19 10.38 1.804 4.339  50  20% P.C. 4285 24 10.17 1.810 4.564  50% I.T.S. 4931 21 9.22 1.777 4.672  50% P.C. 3777 25 13.24 2.035 4.944  50% I.T.S.(R) 4242 25 10.70 1.834 4.609  Species richness  The results from the corrected species richness analysis for the 1995 spring season (Figure 4) shows that compared to control blocks, logging had a positive effect on richness, for both the Mud Lake and Opax Mountain research sites. However, only at the Opax Mountain site were there statistically significant differences in richness values, with 20% patch-cut treatments significantly greater than control treatments. Both Mud Lake and Opax Mountain also had similar richness values, except for 50% LT.S.(R) treatment blocks, where Mud Lake values were somewhat greater. As with the 1995 season, the values for corrected species richness in 1996 (Figure 5) showed an increase in richness in logged treatments when compared to control blocks. Statistically significant differences were found only at the Opax Mountain site, with both 50% patch-cut and 20% I.T.S. treatments significantly greater than 20% patch-cut and control treatments. For both Mud Lake and Opax Mountain, 50% patch-cut treatments contained the greatest species richness. However, both sites did show a substantial drop in species richness compared to 1995 season values (Figure 4). Overall, richness values were fairly similar for both sites, except at 20% removal blocks, where richness values for Mud Lake 20% I.T.S. treatments resembled 20% patch-cut treatments from Opax Mountain, while 20% I.T.S values from Opax Mountain resembled 20% patch-cut treatments from Mud Lake. Corrected species richness values in 1997 (Figure 6) continue to show that logging increased richness in all treatments. Logging also appears to have a greater impact on species richness at the Mud Lake site, where 20% patch-cut, 50% I.T.S. and 50% I.T.S.(R) treatments were substantially greater than their Opax Mountain counterparts.  51  Statistically significant differences in richness were found at both sites, with 50% I.T.S.(R) significantly greater than control and 20% I.T.S. treatments at the Mud Lake site, and 50% patch-cut significantly greater than control, 20% patch-cut, and 50% I.T.S. treatments at the Opax Mountain site. Also, richness values overall rebounded from the low 1996 levels (Figure 5).  52  Control  20%I.T.S.  20%P.C.  50%I.T.S.  1  I  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 4: Corrected species richness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mtn.), spring samples. Rarefaction is used to calculate richness based on a sample size of 10 specimens. F(crit.)=2.40; F(mud)=l .32, F(opax)=3.89 (20%PC ^Control).  Mud L a k e  X  ( s & a & l O p a x Mtn  (0  co <D c .c to  w  I  x  2H  o  <D Q. CO  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 5: Corrected species richness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mtn.), spring samples. Rarefaction is used to calculate richness based on a sample size of 5 specimens. F(crit.)=2.40; F(mud)=1.81, F(opax)=6.36 (50%POControl, 50%PC£20%PC, 20%I.T.S.*Control, 20%I.T.S.*20%PC).  53  6 Mud Lake I  co co <D c  I Opax Mtn  Ix  I  I I  I 7,  X  co O  CJ CD Q. CO  1 H  Control  20%I.T.S.  20%P.C.  50%I.T.S.  50%P.C. 50%I.T.S.(R)  Treatment  Figure 6: Corrected species richness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mtn.), spring samples. Rarefaction is used to calculaterichnessbased on a sample size of 10 specimens. F(crit.)=2.40; F(mud)=3.84 (50%I.T.S.(R)?tControl, 50%I.T.S.(R)?i20%I.T.S.), F(opax)=4.87 (50%PC*Control, 50%PC*20%PC, 50%PC*50% I.T.S.).  54  W h i t t a k e r plots  The Whittaker plots from the 1995 spring season at Mud Lake (Figure 7) show a similar community structure in each of the six experimental treatments, with only a slight increase in the number of intermediate abundant species in the 50% patch-cut treatment. The species rank-abundance model that most closely resembles this community structure is the log-series model. Whittaker plots from the Opax Mountain site (Figure 8) also show a similar community structure for each of the six experimental treatments, with all treatments again resembling the log-series model. Results from the 1996 season at Mud Lake (Figure 9) show plot lines from 20% and 50% patch-cut treatments becoming more even, less steep, with respect to community structure. Control treatment plot lines have become somewhat steeper, compared to 1995 season plots (Figure 7), almost resembling the geometric-series model of community structure. Also, fewer of the more dominant species, those above the 10% abundance mark, were evident in all treatments compared to 1995 (Figure 7). This is especially noticeable in 20% I.T.S. treatments, where one species dominated the carabid community. At the Opax Mountain site (Figure 10) 20% I.T.S. and 50% patch-cut treatments show an increase in numbers of more dominant species, resulting in an almost horizontal plot line just above the 10% abundance mark. Similar to Mud Lake (Figure 9), the control treatments at Opax Mountain have become steeper, reflecting a more dominant community structure, as depicted in the geometric-series model. Results from 1997 Whittaker plots at Mud Lake (Figure 11) show an increase in evenness for all experimental treatments, with control and 20% I.T.S. treatments returning to their 1995 community structure plot lines. Plot lines for 50% I.T.S. and 50%  55  I.T.S.(R) treatments are starting to resemble 20% and 50% patch-cut treatments in appearance, with a more even community structure than seen in previous years. 20% I.T.S. treatments continue to show that one species dominated, similar to the 1996 season (Figure 9). With an increase in the number of intermediate abundant species, control blocks at Opax Mountain returned to a more even community structure of carabid species. 20% I.T.S. and 50% patch-cut treatments have also returned to 1995 levels (Figure 8) in community structure, with fewer abundant species dominating the community. 50% patch-cut treatments have retained their more even, less steep, appearance, as observed in 1996 (Figure 10).  56  Figure 7: Whittaker plots of rank/abundance for Mud Lake 1995 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  Figure 8: Whittaker plots of rank/abundance for Opax Mtn. 1995 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  57  Figure 9 : Whittaker plots of rank/abundance for Mud Lake 1996 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  Figure 10: Whittaker plots of rank/abundance for Opax Mtn. 1996 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  58  Figure 11: Whittaker plots of rank/abundance for Mud Lake 1997 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  Figure 12: Whittaker plots of rank/abundance for Opax Mtn. 1997 season, spring samples. Percent abundance is plotted on a logarithmic scale. For clarity, plot lines are shifted 3 places to the right.  59  Evenness Evenness values for the 1995 spring season (Figure 13) display no obvious trends, other than 50% I.T.S. treatments showing a significantly greater evenness than 50% I.T.S.(R) treatments at Opax Mountain. Results from 1996 (Figure 14) show 50% patch-cut treatments significantly greater than 20% patch-cut and control treatments at Opax Mountain. At the Mud Lake site, the 50% patch-cut treatment increased substantially from 1995 levels (Figure 13), while evenness decreased in the 50% I.T.S. treatment. For both sites, 50% patch-cut treatments show the greatest evenness in the carabid assemblage. Evenness values for the 1997 season (Figure 15) show once again 50% patch-cut treatments significantly greater than 20% patch-cut and control treatments at the Opax Mountain site. A significant difference was also detected at Mud Lake site, with 50% LT.S.(R) significantly greater than 20% I.T.S. treatments. Compared to 1996 levels (Figure 14) at Mud Lake, 50% I.T.S. treatments became more even, while 50% patch-cut and 20% I.T.S. treatments decreased in evenness. The Opax Mountain site was relatively unchanged from the previous year.  60  1.0 I Mud Lake j} Opax Mtn  X  0.8  X  r-i  I  I  0.6  ,)f;  I •  0.4  0.2  ft o.o Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 13: Evenness (Pielou's J') values for 1995 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=1.55, F(opax)=2.76 (50%I.T.S.*50%I.T.S.(R)).  1.0 m%tt Mud Lake EZD Opax Mtn  I  0.8  X  0.6  0.4  0.2  0.0 Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 14: Evenness (Pielou's J') values for 1996 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=0.84, F(opax)=3.38 (50%PC*20%PC, 50%PC*Control).  61  1.0 | Mud Lake j Opax Mtn  0.8  I  I X  I.  0.6 H  0.4 H  0.2 H  0.0 Control  20%I.T.S.  20%P.C.  50%I.T.S.  50%P.C. 50%I.T.S.(R)  Treatment  Figure 15: Evenness (Pielou's J') values for 1997 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=3.28 (50%I.T.S.(R)*20%I.T.S.), F(opax)=3.53 (50%PC*20%PC, 50%PC*Control).  62  Alpha index Alpha index values from the 1995 spring season for Mud Lake and Opax Mountain (Figure 16) showed no statistically significant differences among any of the six experimental treatments. Mud Lake showed higher alpha values for 50% removal treatments, however, no such trend was apparent at Opax Mountain. Control blocks at Mud Lake had the lowest alpha values, while 50% I.T.S. treatments had the highest. At Opax Mountain, all treatments were roughly equal in value, except for the surprisingly high value recorded at the 20% patch-cut treatment block. The 1996 season results (Figure 17) at Opax Mountain show that alpha values for 50% patch-cuts were significantly greater than control treatments. Control blocks at both sites record the lowest alpha values, with 50% I.T.S. treatments at Mud Lake, and 50% patchcut treatments at Opax Mountain recording the highest. Alpha values at Mud Lake substantially increased over 1995 levels (Figure 16) for 20% patch-cut, 50% I.T.S., and 50% patch-cut treatments, with the other treatments remaining relatively unchanged. The only noticeable change at Opax Mountain, compared to 1995 (Figure 16), was the drop in diversity at the 20% patch-cut treatment block. Statistically significant differences in alpha values for the 1997 season (Figure 18) were detected at both sites, with 50% I.T.S.(R) significantly greater than control treatments at Mud Lake, and 50% patch-cuts significantly greater than 50% I.T.S. treatments at Opax Mountain. At Mud Lake there was an increase in the alpha values for 50% LT.S.(R) treatments from 1996 levels (Figure 17), with other treatments remaining at 1996 levels. Opax Mountain showed a small increase in alpha values for 50% patchcut blocks compared to 1996 (Figure 17), but otherwise remained unchanged. Control  63  blocks at Mud Lake, and 50 LT.S. treatments at Opax Mountain recorded the lowest alpha values for 1997.  64  3.5 Mud Lake 3.0  I  I Opax Mtn  I  2.5  ft  2.0 Q. <  I  1.5  1.0  0.5  0.0 Control  20%I.T.S.  20%P.C.  50%I.T.S.  50%P.C  50%I.T.S.(R)  Treatment  Figure 16: Alpha values (+/- s.e.) of the logarithmic series for year 1995 (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=1.34, F(opax)=2.15.  I Mud Lake I Opax Mtn  o. <  3  X  1  H  Control  20%I.T.S.  20%P.C  50%I.T.S.  50%P.C  50%I.T.S.(R)  Treatment  Figure 17: Alpha values (+/- s.e.) of the logarithmic series for year 1996 (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=1.26, F(opax)=3.10 (50%PC^Control).  65  I Mud Lake I Opax Min  «  a. <  I  3 -]  I  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 18: Alpha values (+/- s.e.) of the logarithmic series for year 1997 (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=3.29 (50%I.T.S.(R)*Control), F(opax)=3.26 (50%PC*50%I.T.S.).  66  Shannon-Wiener index Shannon-Wiener diversity indices for the 1995 spring season (Figure 19) show relatively few differences among the six experimental treatments at both Mud Lake and Opax Mountain sites, with only 20% patch-cuts significantly greater than 50% I.T.S.(R) treatments from Opax Mountain. Control blocks at both sites contained the lowest diversity, while 20% I.T.S. treatments at Mud Lake, and 20% patch-cut treatments at Opax Mountain recorded the greatest diversity. In 1996 (Figure 21), differences in diversity between the treatments, compared to the previous season, were more apparent. At Mud Lake, 50% patch-cuts were significantly greater than 50% I.T.S. treatments, while at Opax Mountain, 50% patch-cuts were significantly greater than control, 20% patch-cuts, and 50% I.T.S.(R) treatments, with 20% I.T.S. significantly greater than control, and 20% patch-cut treatments. At the Mud Lake site, control, 20% I.T.S., 50% I.T.S.(R), and 50% I.T.S. treatments all decreased in diversity from 1995 levels (Figure 19), with the other two treatments increasing in diversity only slightly. Opax Mountain showed a slight decrease in almost every treatment, compared to 1995 (Figure 19) levels, with only 50% patch-cut treatments marginally increasing. While control blocks at Opax Mountain were still the lowest in diversity, 50% patch-cut treatments became the most diverse. At Mud Lake, 50% I.T.S. treatments recorded the lowest diversity, while 50% patch-cut treatments recorded the greatest diversity. Results from 1997 (Figure 23) show a slight increase in diversity for each of the six experiment treatments at both sites. Statistically significant differences were also detected at both sites, with 50% I.T.S.(R) and 50% patch-cut treatments significantly  67  greater than control and 20% I.T.S. treatments at Mud Lake, and 50% patch-cuts significantly greater than 20% patch-cut, control, and 50% I.T.S. treatments at the Opax Mountain site. At the Mud Lake site, 50% I.T.S. and 50% I.T.S.(R) treatments show a substantial increase in diversity, rebounding from the low diversity levels in 1996 (Figure 21). The control block at Mud Lake contained the lowest diversity, while 50% I.T.S.(R) treatments contained the highest diversity. At Opax Mountain, 20% patch-cut treatments harbored the least diverse carabid fauna, with 50% patch-cut treatments harboring the greatest.  Simpson's index Simpson's diversity indices for the 1995 spring season (Figure 20) indicate an increase in diversity in all logged treatments, compared to control blocks, at both the Mud Lake and Opax Mountain site, with the only exception being 50% I.T.S.(R) treatments at Opax Mountain, which are slightly less diverse. Statistically significant differences between treatments were detected only at the Opax Mountain site, with 20% patch-cuts significantly greater than 50% I.T.S.(R) treatments. Diversity indices were relatively similar between both sites, except for 50% LT.S.(R) treatments, where the Mud Lake value was considerably greater. Results from the 1996 season (Figure 22) show distinct changes in diversity from the previous season (Figure 20). At Mud Lake, 20% patch-cut, 50% I.T.S. and 50% patchcut treatments had substantially increased in diversity from the previous year, while 20% I.T.S. and 50% I.T.S.(R) treatments had dropped in diversity. At Opax Mountain, 20% I.T.S. and 50% patch-cut treatments had increased in diversity over 1995 levels (Figure 20), with only 20% patch-cut treatments showing a small decline in diversity.  68  Statistically significant differences were detected only at Opax Mountain, with 50% patch-cut and 20% I.T.S. treatments significantly greater than control and 20% patch-cut treatments. Even though noticeable differences in mean diversity were observed at Mud Lake, large amounts of variation cancelled out any significant differences. Control blocks at Opax Mountain, and 20% I.T.S. treatments at Mud Lake, were the least diverse treatments overall, with 50% patch-cut treatments at both sites containing the greatest carabid diversity. The 1997 results for Simpson's diversity indices (Figure 24) show a similar pattern of diversity to 1996 (Figure 22) for all treatments except 50% I.T.S.(R) at Mud Lake, which had rebounded from it's low levels in 1996, to become the most diverse treatment for 1997. For reasons similar to 1996, significant differences were detected only at the Opax Mountain site, with 50% patch-cuts significantly greater than 20% patch-cut, 50% I.T.S. and control blocks.  69  2.0 Mud Lake HggF] Opax Mtn 1.5  1.0  I  I  I  I  I  1 1  A  0.5  0.0 Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 19: Shannon-Wiener Diversity Index values (+/- s.e.) for 1995 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=1.48, F(opax)=2.42 (20%PO50%I.T.S.(R)).  Mud Lske  fHij Opax Mtn  I  I  I  I  T  I  I I  I  3H  11  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 20: Simpson's Diversity Index values (+/- s.e.) for 1995 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=0.88, F(opax)=3.16 (20%PC*50%I.T.S.(R)).  70  2.0 Mud Lake —I  Opax Mtn  1.5 H  X  I  I  i  I.  1.0  0.5  0.0 Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 21: Shannon-Wiener Diversity Index values (+/- s.e.) for 1996 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=3.25 (50%PG#50%I.T.S.), F(opax)=6.00 (50%POControl, 5 0 % P C * 2 0 % P C , 50%PCV50%I.T.S.(R), 20%I.T.S.#Control, 20%I.T.S.*20%PC).  7  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 22: Simpson's Diversity Index values (+/- s.e.) for 1996 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=1.13, F(opax)=5.82 (50%PC*Control, 5 0 % P C * 2 0 % P C , 20%I.T.S.*Control, 20%I.T.S.*20%PC).  71  2.0  ix  m§tm Mud Lake f » ) Opax Mtn  I  1.5  I  i  I  I  X  X  1.0  0.5  0.0 Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 23: Shannon-Wiener Diversity Index values (+/- s.e.) for 1997 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=4.40 (50%I.T.S.(R)*Control, 50%I.T.S.(R)*=20%I.T.S., 5 0 % P C * C o n t r o l , 50%PC*20%I.T.S.), F(opax)=4.86 ( 5 0 % P C * 2 0 % P C , 5 0 % P C * C o n t r o l , 50%PC*50%I.T.S.).  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 24: Simpson's Diversity Index values (+/- s.e.) for 1997 season (Mud Lake and Opax Mtn.), spring samples. F(crit.)=2.40; F(mud)=2.15, F(opax)=4.82 ( 5 0 % P C * 2 0 % P C , 50%PC#50%I.T.S., 50%PC^Control).  72  T a x o n o m i c distinctness  Taxonomic distinctness (TD) values for the 1995 spring season (Figure 25) show relatively small differences in distinctness between treatments at Mud Lake or Opax Mountain. Control blocks from both sites had the highest distinctness, with 50% I.T.S.(R) treatments at Mud Lake and 50% I.T.S. treatments at Opax Mountain recording the lowest distinctness. Statistically significant differences were detected only at the Opax Mountain site, with control, 50% patch-cut, and 20% I.T.S. treatments significantly greater than 50% I.T.S. treatments. Except for 50% I.T.S. treatments, each of the six treatments at Opax Mountain were slightly more distinct than their counterpart treatments from Mud Lake. Results from 1996 for T D (Figure 26) show virtually no changes at the Opax Mountain site from 1995 levels (Figure 25), while at Mud Lake a slight drop in distinctness for the 20% I.T.S. and control blocks was observed, with a more substantial drop in the 50% I.T.S. treatment. At the Mud Lake site, 50% patch-cut treatments in 1996 contained the most distinct carabid community, with 50% I.T.S. treatments containing the least distinct community. Similar to 1995 results (Figure 25), only the Opax Mountain site showed statistically significant results, with control, 50% patch-cut, and 50% LT.S.(R) treatments significantly greater than 50% I.T.S. treatments. In 1997 (Figure 27), T D values had returned to 1995 levels (Figure 25) at Mud Lake. At Opax Mountain, distinctness remained unchanged from 1996 levels (Figure 26). Control, 50% I.T.S.(R), and 50% patch-cut treatments were once again significantly greater in T D than 50% I.T.S. treatments, at the Opax Mountain site. With the exception  73  of 50% I.T.S. treatments, Opax Mountain treatments were once again slightly more distinct than their Mud Lake counterparts.  74  B  100  I  80  60  Mud Lake Opax Mtn  I A  40  20  r  Control  20%I.T.S.  20%P.C  50%I.T.S.  L_J  50%P.C  50%I.T.S.(R)  Treatment  Figure 25: Taxonomic distinctness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=1.42, F(opax)=3.90 (Control, 50%PC, 20%I.T.S.*50%I.T.S.).  100 Mud La j Opax Mtn Ke  [•  80  X  I  I  X 60  40  A  20  Control  20%I.T.S.  20%P.C.  50%I.T.S.  50%P.C.  50%I.T.S.(R)  Treatment  Figure 26: Taxonomic distinctness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=0.98, F(opax)=4.14 (Control, 50%PC, 50%I.T.S.(R)jt50%I.T.S.).  75  ioo H  H I  80  Mud Lake j Opax Mtn  I  H  60  40 i  20  H  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 27: Taxonomic distinctness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=0.69, F(opax)=3.30 (Control, 50%I.T.S.(R), 50%PC*50%I.T.S.).  76  Average taxonomic distinctness  Average taxonomic distinctness (aveTD) graphs from the 1995 spring season (Figure 28) show very similar values across all treatments for both sites, with the exception of a lower average distinctness at 50% I.T.S. treatments, and a higher aveTD in 50% patchcut treatments at Mud Lake, compared to control block values. Statistically significant differences were detected only at the Mud Lake site, with 50% patch-cuts significantly greater than 50% I.T.S treatments. Results from 1996 (Figure 29) show virtually no change for Opax Mountain treatments from the previous year, while at Mud Lake, all treatments showed a slight decrease in aveTD, with 50% I.T.S. treatments showing a much larger decrease. In 1997 (Figure 30) aveTD values for Mud Lake treatments returned to 1995 levels (Figure 28), with only patch-cut treatments slightly lower in aveTD than in 1995. While at Opax Mountain in 1997, treatments remain unchanged in aveTD from previous years. Results from the funnel graphs for the three-year period (Figures 34, 36, and 38), show all treatments falling inside the expected 95% confidence intervals for aveTD.  77  Mud Lake  100  I v ^ Opax Mtn  80  P.  CD > CO  X  60 -\  40  20 A  Control  20%I.T.S.  20%P.C  50%I.T.S.  50%P.C 50%I.T.S.(R)  Treatment  Figure 28: Average taxonomic distinctness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=3.24 (50%PC*50%I.T.S.), F(opax)=1.31.  100  I  Mud Lake I Opax Mtn  80  P  I  60  >  CO 40 A  20  Control  20%I.T.S.  20%P.C  50%I.T.S.  50%P.C 50%I.T.S.(R)  Treatment  Figure 29: Average taxonomic distinctness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=2.04, F(opax)=1.04.  78  100  Mud Lake ImM Opax Mtn  80  P_  60  CD > CO  40 H  20 H  Control  20%I.T.S.  20%P.C  50%I.T.S.  50%P.C 50%I.T.S.(R)  Treatment  Figure 30: Average taxonomic distinctness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=2.31, F(opax)=l .82.  79  V a r i a t i o n i n taxonomic distinctness  Variation in taxonomic distinctness (varTD) values for the 1995 spring season (Figure 31) show relatively minor differences between treatments at Opax Mountain, while differences between treatments at Mud Lake are much more apparent. The 20% patchcut, 50% I.T.S. and 50% I.T.S.(R) treatments at Mud Lake were substantially higher than other treatments. However, only 50% I.T.S. treatments showed a significantly greater varTD than 50% patch-cut treatments. In both sites the 50% patch-cut treatments recorded the lowest varTD, with 50% I.T.S. treatments recording the highest value. In 1996 (Figure 32), the Mud Lake site showed a decrease in varTD values at both control and 50% I.T.S. treatments, with all other treatments, except for the 50% I.T.S.(R) treatment, showing an increase. At Opax Mountain, each of the six treatments increased in varTD from 1995 levels (Figure 31), resulting in less noticeable differences between treatments overall. Statistically significant differences were observed only at the Mud Lake site, with 20% patch-cut treatments significantly greater than control blocks. Results in 1997 (Figure 33) show that control and 50% I.T.S. treatments at Mud Lake were returning to almost 1995 levels (Figure 31), with other treatments showing a slight decrease in varTD from the previous year (Figure 32). At Opax Mountain, each treatment had decreased in varTD from 1996 (Figure 32), and this was especially so for 20% and 50% patch-cut treatments. Although statistically significant differences were detected at the Opax Mountain site, the Tukey test was unable to detect which treatment differed from which.  80  As with aveTD, the funnel graphs for varTD for the three-year period (Figures 35, 37, and 39), failed to show any treatments falling outside of the 95% confidence intervals, at either site.  81  1000 Mud Lake  Q  800  A  600  A  w*w»i Opax Mtn  I  I r—n  X  CO >  I  400  200  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 31: Variation in taxonomic distinctness values (+/- s.e.) for 1995 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=2.99 (50%I.T.S.#50%PC), F(opax)=1.18.  1000  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 32: Variation in taxonomic distinctness values (+/- s.e.) for 1996 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=3.33 (20%PC*Control), F(opax)=0.76.  82  800 I Mud Lake  700 -  I  ] Opax Mtn  600 -  I  500  i 400  fi  300  200  H  100  0 Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 33: Variation in taxonomic distinctness values (+/- s.e.) for 1997 season (Mud Lake and Opax Mountain), spring samples. F(crit)=2.40; F(mud)=1.48, F(opax)=2.54.  83  90-TCON(0)  5OPC{OT  A 85-{-  S0rTS(O)  20PG(M) ±  2 0 ,  ^  ( P )  20PC(O) 20ITS(M) A  GON(M)i  ' 50R(M)  sorrs(M)  '50PJ3(M)  80-4-  i  70-4-  65 " 12  18  18  14  22  20  Number of species  Figure 34: The 95% 'probability funnel' for average taxonomic distinctness, with 1995 spring sample aveTD values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  800-r-  700r4r CON(M) eorrs(O) 800-4^  5  . 500-4-  20PC(M)  2on%(M)  20rrs(O)  *  A "  50PC(M)  2  0  ^  {  Q  )  S0PC(O)  A "" ' 50R(O)  400-4-  300 12  ;14  20  18  22;  Number of species:  Figure 35: The 95% 'probability funnel' for variation in taxonomic distinctness, with 1995 spring sample varTD values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  84  100"!"'  ,90-+S0R(O)  20IT5(o) 50ITS(O)  60PC(O)  20nfc(M) CON(M)  20PC(M) 80-4-  S0R(M):  A-'  'SOP^M)  sorrsfM)  Figure 36: The 95% 'probability funnel' for average taxonomic distinctness, with 1996 spring sample aveTD values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  1000-r-  8004-  5orrs(0)  CON(M)  20PC(M)  •a  S0R(M)  2orrs(M)  600-4„$0RCQ)  50l4(M)  ^2orfl(Crj— S(M>501T  CON(Or  50PC(O) 20PC(O)  400-+-  200 10  12  141  16  Number of species  Figure 37: The 95% 'probability funnel' for variation in taxonomic distinctness, with 1996 spring sample varTD values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  85  95-r-  80-4CON(M) A  20ITS{O)  SOITS(O)  85-4-  20ITS(M)  20PC(O) A  5 0  80-4-  ^°)  ^0PC(M)_ 50R(M)  BOPCKM)  5orrs(M)  754-  704-  854-  60  15'  io;  20  25  Number of species  Figure 38: T h e 95% 'probability funnel' for average taxonomic distinctness, with 1997 spring sample a v e T D values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  900-r-  aoo4-  50rrs(M)  50PC(M).  20PC(M)  6004-  A_50Rfl4)..  50R(O>—  20PC(O)A  2orrs(0)  5004• CGN(M) • CON(0)  :ji 5orrs(0)  2orrs<M) ^ 50PC(G)  ' Jk  Number of species  Figure 39: T h e 95% 'probability funnel' for variation in taxonomic distinctness, with 1997 spring sample varTD values (Mud Lake and Opax Mountain), derived from 1000 independent simulations for each subset size drawn randomly from the 58 Opax area carabid species.  86  Cluster analysis Cluster analysis for the 1995 spring season (Figure 40) shows Mud Lake and Opax Mountain clustering separately, with the exception of control and 20% I.T.S. treatments at Mud Lake, which cluster together with Opax treatments. The higher percent similarities between treatments at Opax Mountain shows a much more similar fauna, than between treatments at the Mud Lake site. At Opax Mountain, the two patch-cut treatments cluster together, as well as the two I.T.S. treatments, with 50% I.T.S.(R) from Opax Mountain clustering closer to 20% I.T.S. at Mud Lake. Control blocks from both sites cluster closer to I.T.S. than to patch-cut treatments. Overall, the treatments from both sites were at least 70.2 % similar to each other. In 1996 (Figure 42), cluster analysis shows a complete separation between Mud Lake and Opax Mountain sites. A higher percent similarity between treatments at Opax Mountain continued to show that these treatments are less affected by the logging practices, than the same treatments are at Mud Lake. At both sites, patch-cut treatments cluster together, reflecting their similar carabid fauna. Control blocks at Mud Lake cluster closer to 20% I.T.S. treatments, while at Opax Mountain, control blocks cluster with 50% I.T.S.(R) treatments. A 63.2% similarity between all treatments in both sites was observed for the 1996 season. Results from the 1997 season (Figure 44) show Mud Lake and Opax Mountain once again clustering separately, with the exception of the control blocks from Mud Lake, which cluster closer to control and I.T.S. treatments from Opax Mountain. At Mud Lake, patch-cut treatments once again cluster together, with 20% I.T.S. treatments clustering with 50% LT.S.(R), and 50% I.T.S. clustering with patch-cut treatments. At the Opax  87  Mountain site, 20% patch-cut treatments cluster with 50% LT.S.(R) treatments, while 50% patch-cuts are the next most similar fauna. Control blocks at Opax Mountain cluster with 20% I.T.S. treatments, with 50% I.T.S. treatments clustering next to these. Overall, a 66.2% similarity was observed between all treatments at both sites, up from the previous years 63.2% level.  M u l t i d i m e n s i o n a l scaling ( M D S )  The M D S graph from the 1995 spring season (Figure 41) corroborates the results from cluster analysis in 1995 (Figure 40). In M D S analysis, the different treatments within each site appear closer to each other on the graph, than to the similar treatments at the other site. The exception to this is with the 20% I.T.S. and control treatments from Mud Lake, which drift closer to Opax Mountain treatments on the map, than to the other treatments at Mud Lake. As with the higher percentages in similarity shown in the cluster analysis for the Opax Mountain site treatments, the results from the MDS graph show these treatments in closer proximity, which reflects the great similarity in carabid faunas between treatments. At both sites, patch-cut treatments plot closer to each other than to similar percent removal treatments harvested by individual-tree selection. Control treatments at Opax Mountain show a similar carabid assemblage to 50% I.T.S. treatments, by their close proximity. In 1996 (Figure 43), results from the M D S graph show an even greater separation in distance between Mud Lake and Opax Mountain, than what was mapped in the previous year (Figure 41). Greater similarity is once again seen at Opax Mountain, compared to the more spread out treatments observed at Mud Lake. Mud Lake shows patch-cut  88  treatments in close proximity to 50% I.T.S. treatments, with 20% I.T.S., 50% I.T.S.(R) and control treatments grouping amongst themselves. Both sites show patch-cut treatments in close proximity. Control treatments from Mud Lake plot closer to 20% I.T.S. treatments, while at Opax Mountain they plot closer to 50% LT.S.(R) treatments. Results from 1997 (Figure 45) continue to show a separation of both sites, with each treatment showing a more similar carabid fauna to its surroundings than to the similar harvested treatment from the other site. Similar to 1995 (Figure 41) results, control blocks drift closer to the Opax Mountain fauna in similarity, than to its own Mud Lake fauna, with 20% I.T.S. treatments not far behind. Patch-cut treatments from Mud Lake show a closeness reflecting a similar fauna, while at Opax Mountain 20% patch-cuts plot closer to 50% I.T.S.(R) treatments, with 50% patch-cuts next closest to these. M D S results at Opax Mountain also showed a similar carabid assemblage between control and 20% I.T.S. treatments.  89  50%P.C.(M) 20%P.C.(M) 81.2  77.7  50%I.T.S.-R(M)  75.1  50%I.T.S.(M) Control(M) 70.2  20%I.T.S.(M) 85.0  50%I.T.S.-R(O)  81.3  20%P.C.(OJ_ 50%P.C.(O_^_ 20%I.T.S.(O) 50%I.T.S.(O)_ Control(O)  76.9  83.8 78.7  88.0 84.4  Figure 40: Dendrogram for hierchical clustering of treatment data from Mud Lake and Opax Mtn. sites (1995), spring samples, using group-average linking of Bray-Curtis similarities calculated on 4th root-transformed abundance data. Note: M=Mud Lake, 0=Opax Mtn.  Figure 41: MDS of species abundances on treatment data for Mud Lake and Opax Mtn.(1995), spr. samples. 1/7=20%P.C.(M/O), 2/8=20%I.T.S.(M/O), 3/9=50%P.C.(M/O), 4/10=50%I.T.S.-R(M/O), 5/11=50%I.T.S.(M/O), 6/12=Control(M/0), (Stress=0.09). Note: M=Mud Lake, 0=Opax Mtn.  90  20%P.C.(M)  77.9  50%P.C.(M)  73.5 50%I.T.S.(M) 50%I.T.S.-R(M)  69.8  20%I.T.S.(M)_  72.9 83.9  Control(M) 20%I.T.S.(O)  63.2  20%P.C.(O) 50%P.C.(O)  81.8 84.9 77.6  50%I.T.S.(O) 50%I.T.S.-R(O) Control(O)  83.5 85.4  Figure 42: Dendrogram for hierchical clustering of treatment data from Mud Lake and Opax Mtn. sites (1996), spring samples, using group-average linking of Bray-Curtis similarities calculated on 4th root-transformed abundance data. Note: M=Mud Lake, 0=Opax Mtn.  Figure 43: MDS of species abundances on treatment data for Mud Lake and Opax Mtn. (1996), spr. samples. 1/7=20%P.C.(M/O), 2/8=20%I.T.S.(M/O), 3/9=50%P.C.(M/O), 4/10=50%I.T.S.-R(M/O), 5/11=50%I.T.S.(M/O), 6/12=Control(M/0), (Stress=0.1). Note: M=Mud Lake, 0=Opax Mtn.  91  20%I.T.S.(M)  T73  50%I.T.S.-R(M) 20%P.C.(M)  72.4  83.1  50%P.C.(M)  76.9  50%I.T.S.(M) 50%P.C.(O)  66.2  20%P.C.(O)  79.8 86.1  50%I.T.S.-R(O)  78.1  Control(M) 50%I.T.S.(O)  81.2  20%I.T.S.(O}_ Control(O)  84.6 87.6  Figure 44: Dendrogram for heirchical clustering of treatment data from Mud Lake and Opax Mtn.(1997) sites, spring samples, using group-average linking of Bray-Curtis similarities calculated on 4th root-transformed abundance data. Note: M=Mud Lake, 0=Opax Mtn.  Figure 45: MDS of species abundances on treatment data for Mud Lake and Opax Mtn.(1997), spr. samples. 1/7=20%P.C.(M/O), 2/8=20%I.T.S.(M/O), 3/9=50%P.C.(M/O), 4/10=50%I.T.S.-R(M/O), 5/11=50%I.T.S.(M/O), 6/12=Control(M/0), (Stress=0.07). Note: M=Mud Lake, 0=Opax Mtn.  92  Dominance structure Results from spring samples at Mud Lake (Table 11) show that the top four most abundant carabid species from control blocks, remained the most abundant four species in the five harvested treatments, but only switched in places of dominance and percent abundance. In terms of dominance position only, the closest resemblance to the carabid community dominance structure observed in control blocks was found in 20% I.T.S. treatments. Looking at single species changes, Pterostichus adstrictus became the most dominant species in 50% P.C. and 50% I.T.S. treatments, Cymindis unicolor was more dominant in control than in harvested blocks, and both Bembidion dyschirinum and Notiophilus directus had the highest percent abundance in patch-cut treatments. Results from entire year samples at Mud Lake (Table 12) show that the top four most abundant carabid species from control blocks, remained the most abundant four species in the five harvested treatments, but only switched in places of dominance and percent abundance. The closest resemblance to the carabid community dominance structure observed in control blocks was found in 20% I.T.S. treatments. For single species changes, Pterostichus adstrictus became one of the dominant two species in 50% P.C. and 50% I.T.S.(R) treatments, while Scaphinotus angusticollis dropped out of the ten most dominant list, except in 20% I.T.S. treatments, where it was slightly more dominant than what was observed from control blocks. Results from spring samples at Opax Mountain (Table 13) show that the top five most abundant carabid species from control blocks, remained the most abundant five species in the five harvested treatments, but only switched in places of dominance and percent abundance. In terms of dominance position only, 20% patch-cut treatments most closely  93  resembled the carabid community dominance structure observed in control blocks. Looking at single species changes, Scaphinotus angusticollis dropped out of the top ten most abundant species list in all harvested treatments, while Bembidion dyschirinum increased it's dominance position, as well as it's percent abundance in 50% patch-cut treatments. Results from entire year samples at Opax Mountain (Table 14) show that the top four most abundant carabid species from control blocks, remained the most abundant four species in the five harvested treatments, but only switched in places of dominance and percent abundance. The closest resemblance to the carabid community dominance structure observed in control blocks was found in 20% I.T.S. treatments. For single species changes, Scaphinotus angusticollis dropped out of the top ten most abundant species list in all harvested treatments, except for 50% I.T.S. and 20% patch-cut treatments, Bembidion dyschirinum increased it's dominance position, as well as it's percent abundance in 50% patch-cut treatments, while Pterostichus adstrictus had a higher dominance position in every harvested treatment, compared to control blocks.  94  Table 11: Dominance position (and percent abundance) of the major species in the experimental treatments at the Mud Lake site, based on spring samples, 1995-1997.  Pterostichus neobrunneus Carabus taedatus agassii Pterostichus adstrictus Calathus ingratus Calathus advena Scaphinotus marginatus Cymindis unicolor Bembidion dyschirinum Syntomus americanus Notiophilus directus  Control 1. (34.32) 2. (22.13) 3. (17.08) 4. (16.12) 5. (5.53) 6. (1.76) 7. (1.20) 8. (0.40) T9. (0.24) T9. (0.24)  20%I.T.S. 1. (42.65) 2. (15.64) 3. (13.94) 4 (12.19) 5. (8.29) 6. (2.07) 8. (0.82) 10.(0.63) 9. (0.75) 7. (0.88)  20%P.C. . 1. (28.45) 3. (14.83) 2. (20.77) 4. (12.65) 9. (1.92) T10. (0.87) T10. (0.87) 7. (3.49) T14. (0.52) 6. (4.54)  50%I.T.S. 1. (28.49) 2. (23.98) 3. (21.51) 4. (11.05) 18. (0.29) T15. (0.44) T11. (0.87) 6. (1.74) 14. (0.58) 7. (1.45)  50%P.C. 2. (26.72) 4. (9.60) 1. (29.12) 3. (10.05) 6. (3.70) 10. (1.82) 11. (1.30) 7. (3.11) 13. (0.65) 8. (2.85)  50%I.T.S.(R) 2. (24.67) 3. (11.33) 1. (38.12) 4. (7.14) 6. (3.57) 9. (1.24) 10. (1.16) 11. (1.09) 15. (0.39) 7. (2.17)  Table 12: Dominance position (and percent abundance) of the major species in the experimental treatments at the Mud lake site, based on complete year samples, 1993-1997.  Carabus taedatus agassii Pterostichus neobrunneus Calathus ingratus Pterostichus adstrictus Scaphinotus marginatus Synuchus impunctatus Calathus advena Scaphinotus angusticollis Cymindis unicolor Cymindis cribicollis  Control 1. (31.21) 2. (20.01) 3. (16.95) T4. (8.13) T4. (8.13) 6. (5.92) 7. (4.51) 8.(1.69) .. 9. (1.58) 10. (0.62)  20%I.T.S. 2. (24.48) 1. (26.63) 3. (11.10) 5. (7.61) 4. (7.64) 8. (1.90) 6. (7.02) 7. (3.12) 9. (1.71) 11. (0.87)  20%P.C. 1. (27.86) 2. (17.31) 3. (14.33) 4. (11.67) 5. (6.60) 6. (5.84) 9. (2.05) 14. (0.72) 13.(0.81) T19. (0.24)  50%I.T.S. 1. (37.24) 2. (17.38) 3. (14.83) 4. (12.22) 5. (3.76) 7.(1.72) 16. (0.45) T11. (0.89) 13. (0.83) 8. (1.34)  50%P.C. 2. (20.86) 3. (19.56) 4. (12.40) 1. (20.89) 5. (4.71) T7. (2.92) 6. (4.08) 15. (0.56) 11. (1.62) T19. (0.20)  50%I.T.S.(R) 1. (24.94) 3. (15.94) 4. (14.79) 2. (23.45) 6. (3.18) 8. (2.41) 5. (4.37) 16. (0.46) 9.(1.34) T13. (0.61)  Table 13: Dominance position (and percent abundance) of the major species in the experimental treatments at the Opax Mountain site, based on spring samples, 1995-1997.  Pterostichus neobrunneus Carabus taedatus agassii Calathus advena Calathus ingratus Pterostichus adstrictus Notiophilus directus Scaphinotus marginatus Scaphinotus angusticollis Cymindis unicolor Bembidion dyschirinum  Control 1. (36.11) 2. (29.23) 3. (22.88) 4. (4.54) 5. (2.13) 6. (1.53) 7. (1.33) 8. (0.64) 9. (0.60) 10. (0.44)  20%I.T.S. 2. (27.97) 1. (29.61) 4. (15.12) 3. (16.90) 5. (3.78) 7. (1.69) 6. (1.63) T14. (0.04) 8. (0.80) 9. (0.71)  20%P.C. 1. (41.42) 2. (18.54) 3. (17.35) 4. (9.87) 5. (5.94) 6. (1.87) 7. (1.23) T16. (0.07) 8. (1.01) 9. (0.82)  50%I.T.S. 1. (41.93) 3. (15.39) 2. (19.84) 4. (9.79) 5. (9.04) T6. (1.04) T6. (1.04) 11. (0.16) 8. (0.88) 10. (0.23)  50%P.C. 2. (23.99) 1. (27.90) 4. (10.45) 3. (13.39) 5. (9.29) 6. (3.73) 8. (1.26) T17. (0.09) T10. (0.93) 7. (3.64)  50%I.T.S.(R) 2. (26.24) 3. (21.65) 1. (32.07) 4. (6.58) 5. (5.68) T7. (1.02) 6. (2.99) T11. (0.21) T7. (1.02) 9. (0.85)  Table 14: Dominance position (and percent abundance) of the major species in the experimental treatments at the Opax Mountain site, based on complete year samples, 1993-1997.  Carabus taedatus agassii Pterostichus neobrunneus Calathus advena Calathus ingratus Scaphinotus marginatus Notiophilus directus Scaphinotus angusticollis Pterostichus adstrictus Cymindis unicolor Bembidion dyschirinum  Control 1. (40.71) 2. (24.84) 3. (18.21) 4. (5.19) 5. (2.84) 6. (2.47) 7.(1.97) 8. (1.53) 9. (0.96) 10. (0.39)  20%I.T.S. 1. (39.17) 2. (18.51) 4. (11.28) 3. (16.32) 5. (3.77) 6. (3.64) T14. (0.20) 7. (2.39) 8. (1.48) 10. (0.54)  20%P.C. 2. (28.40) 1. (31.58) 3. (16.10) 4. (9.52) 8. (1.63) 6. (2.92) 10. (0.40) 5. (4.78) 7. (2.10) 9. (0.77)  50%I.T.S. 2. (25.43) 1. (31.29) 3. (18.80) 4. (10.30) 6. (1.97) 7. (1.64) 9. (1.28) 5. (6.79) 8. (1.32) 12. (0.18)  50%P.C. 1. (37.83) 2. (16.34) 4. (8.68) 3. (12.66) 8. (2.20) 6. (4.37) 17. (0.32) 5. (7.23) 9. (1.83) 7. (2.59)  50%I.T.S.(R) 1. (31.64) 3. (19.17) 2. (26.76) 4. (6.39) 5. (4.50) 7. (1.93) 12. (0.47) 6. (4.31) 8. (1.18) T10. (0.66)  Species analysis Pterostichus neobrunneus The most abundant species overall in the spring samples was P. neobrunneus (Table 4). P. neobrunneus was more abundant at the Opax Mountain site, with 5,057 specimens captured (Table 6), compared to only 2,359 specimens from Mud Lake (Table 5). Although P. neobrunneus was most numerous in 20% I.T.S. treatments at Mud Lake, and 50 I.T.S. treatments at Opax Mountain (figure 46), it was also found in relatively high numbers in other treatments. The results from the non-parametric A N O V A test indicated a statistically significant difference between every treatment in both sites.  Carabus taedatus agassii The second most abundant species collected in spring samples was C. taedatus agassii (Table 4). C. taedatus agassii was also more abundant at the Opax Mountain site, with 3,467 specimens (Table 6), compared to 1,154 from Mud Lake (Table 5). Both sites showed a similar response to logging (Figure 47), with the least disturbed control and 20% I.T.S. treatments harboring the most specimens, while patch-cut treatments and higher percent removal blocks contained the fewest. No significant differences were detected between any of the treatments at Mud Lake, or between 20% patch-cut, 50% patch-cut, and 50% I.T.S. treatments at Opax Mountain.  Calathus advena The third most abundant species in spring samples was C. advena (Table 4). Similar to Pterostichus neobrunneus and Carabus taedatus agassii, Calathus advena was  97  considerably more abundant at the Opax Mountain site, with 2,958 collected (Table 6), compared to only 328 at Mud Lake (Table 5). As with Pterostichus neobrunneus (Figure 46), no recognizable pattern of higher captures was observed for the Mud Lake population of Calathus advena. However, the Opax Mountain population did show a slight higher capture rate for 50% I.T.S.(R) treatments, though not a statistically significant one (Figure 48). Kruskal-Wallis tests showed a non-significant difference between Control and 20% I.T.S. treatments at Mud Lake, and between 50% IT.S(R) and 20% I.T.S. treatments at Opax Mountain.  Calathus ingratus The fifth most abundant species collected in the spring samples between the years 1995-1997 was C. ingratus (Table 4). Of the total 2,362 specimens, 1,499 individuals were caught at the Opax Mountain site (Table 6), while 863 were caught from the Mud Lake site (Table 5). The results from the Mud Lake population of C. ingratus showed a significantly higher occurrence for the less disturbed treatments (Figure 49), while the Opax Mountain population appeared to be more common in the I.T.S. treatments, though only significantly for the 20% I.T.S. treatment. Results from Kruskal-Wallis tests show non-significant differences between 50% and 20% patch-cut treatments at Mud Lake, and between 50% I.T.S. and 50% patch-cut treatments at Opax Mountain.  Pterostichus adstrictus Spring samples yielded a total of 2,671 specimens of P. adstrictus between the years 1995-1997, making it the fourth most abundant species overall (Table 4). Unlike the  98  other four abundant species, P. adstrictus was more dominant at the Mud Lake site, with 1,764 specimens collected (Table 5), compared to 907 from Opax Mountain (Table 6). While the Opax Mountain population of P. adstrictus appeared to occur more abundantly in the more disturbed 50% removal treatments, the Mud Lake population occurred more abundantly in 50% I.T.S.(R) treatments over all other treatments (Figure 50). KruskalWallis analysis detected no statistically significant differences between 50% I.T.S.(R) and 50% patch-cut treatments, or between control and 50% I.T.S. treatments at Mud Lake, nor between the 50% patch-cut and 50% I.T.S.(R) treatments at Opax Mountain.  Scaphinotus marginatus Although nowhere as dominant as the previous five species, S. marginatus did amount to over 1.5% of the total catch (Table 4), with 234 beetles collected from Opax Mountain (Table 6), and 112 from Mud Lake (Table 5). At Opax Mountain S. marginatus was more common in 50% I.T.S.(R) treatments than other treatments, while at Mud Lake there was no recognizable difference for any percent removal (20% or 50%) or method of harvest (I.T.S., I.T.S.(R), or patch-cut) (Figure 51). Kruskal-Wallis analysis failed to detect a significant difference between any of the treatments at Opax Mountain, while at Mud Lake, 20% I.T.S. and 50% patch-cut treatments, as well as 20% I.T.S. and control treatments showed a non-significant difference in abundance.  99  Bembidion dyschirinum Accounting for almost 1.3% of the total catch of carabid beetles from spring samples (Table 4), B. dyschirinum was one of the few species to clearly show a recognizable correlation with method of harvest. At both Mud Lake and Opax Mountain, a significantly greater number of beetles were caught in patch-cut treatments, compared to other treatments in either site (Figure 52). This suggests an obvious preference of B. dyschirinum for disturbed, open habitats. Results from Kruskal-Wallis failed to detect a significant difference 50% I.T.S.(R) and 50% I.T.S. treatments at Mud Lake, or between 50% LT.S.(R) and 20% I.T.S. treatments at Opax Mountain.  Notiophilus directus Similar to all but two of the dominant carabid species, N. directus was clearly most common in the Opax Mountain site habitat, with 282 (Table 6) of the total 302 specimens (Table 4). At Opax Mountain, N. directus occurred more in patch-cut treatments than in control or I.T.S. treatments, while at Mud Lake abundance patterns were more difficult to detect, owing to the low number of beetles collected from the site (Figure 53). No significant differences were detected in any of the treatments from either site, using the non-parametric Kruskal-Wallis analysis.  Cymindis unicolor Just under two-thirds of the species C. unicolor were collected from the Opax Mountain site (Table 6), showing that this species was more dominant here than at Mud Lake (Table 5). Although the species at Opax Mountain was captured more often in logged  100  treatments than in control blocks, no such differences were observed for either percent removal or method of harvest at the Mud Lake site (Figure 54). The non-parametric Kruskal-Wallis analysis failed to detect any significant differences between any of the treatments in either site.  Syntomus americanus In addition to Pterostichus adstrictus, Syntomus americanus was the only other dominant carabid species that was more frequently collected at Mud Lake than at Opax Mountain, with over three-quarters of beetles caught at the Mud Lake site (Table 5). Similar to the species Bembidion dyschirinum (Figure 52), Syntomus americanus was most common in patch-cut treatments at both sites (Figure 55). Non-parametric A N O V A results failed to detect a significant difference between 20% and 50% I.T.S. treatments at Mud Lake, or between 20% I.T.S. and 50% I.T.S.(R), or 50% I.T.S. and control treatments at Opax Mountain.  101  Figure 46: Mean  specimen number (+/- s.e.) of P. neobrunneus for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=11.07; chi-square(mud)=31.33, chi-square(opax)=36.89.  5A Q. CD  53  CL  HI  Mud Lake  [lip  Opax Mtn  T 4 T  C  CD  £ o  CD Q. CO  C  CO CD  Control  L Ii i i  20%I.T.S. 20%P.C.  50%I.T.S. 50%P.C. 50%I.T.S.(R)  Treatment  Figure 47: Mean specimen number (+/- s.e.) of C. taedatus agassii for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=l 1.07; chi-square(mud)=3.74 (Control=20%I.T.S.=20%P.C.=50%I.T.S.=50%P.C.=50%I.T.S.(R)), chi-square(opax)=12.22 (50%P.C.=20%P.C.=50%I.T.S.).  102  Mud Lake MEM Opax Mtn Q. CO  I  CD  Q. * C CD  E CJ CD  I  Q. CO  c CO CD  a si  1  Control  20%I.T.S.  -X.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 48: Mean specimen number (+/- s.e.) of C. advena for seasons 1995-1997 (Mud Lake and Opax Mm.), based on spring samples only. Chi-square(crit.)=11.07; chi-square(mud)=27.44 (20%I.T.S.=Control), chi-square(opax)=19.10 (50%I.T.S.(R)=20%I.T.S.).  Mud Lake  X  pg] Opax Mtn  Q. CO  CD Q.  2A  C CD  E o CD Q. CO  14  CO CD  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 49: Mean specimen number (+/- s.e.) of C. ingratus for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=l 1.07; chi-square(mud)=13.17 (50%P.C.=20%P.C), chi-square(opax)=44.72 (50%I.T.S.=50%P.C).  103  4  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 50: Mean specimen number (+/- s.e.) of P.adstrictus for seasons 1995-1997 (Mud Lake and Opax Mm.), based on spring samples only. Chi-square(crit.)=11.07; chi-square(mud)=26.38 (50%I.T.S.(R)=50%P.C, Control=50%I.T.S.), chi-square(opax)=20.82 (50%P.C.=50%I.T.S.(R)).  0.6  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 51: Mean specimen number (+/- s.e.) of S. marginatus for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=l 1.07; chi-square(mud)=15.28 (50%P.C.=20%I.T.S., 20%I.T.S.=Control), chi-square(opax)=8.30 (50%I.T.S.(R)=20%I.T.S.=Control=20%P.C.=50%I.T.S.=50%P.C).  104  0.6  Control  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 52: Mean specimen number (+/- s.e.) of B. dyschirinum for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=11.07; chi-square(mud)=26.17 (50%I.T.S.(R)=50%I.T.S.), chisquare(opax)=29.00(50%I.T.S.(R)=20%I.T.S.).  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure S3: Mean specimen number (+/- s.e.) of N. directus for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=l 1.07; chi-square(mud)=5.03 (20%I.T.S.=50%P.C.=20%P.C.=50%I.T.S.(R)=50%I.T.S.=Control), chi-square(opax)= 10.95 (50%P.C.=20%P.C.=20%I.T.S.=Control=50%I.T.S.=50%I.T.S.(R)).  105  0.25 H I M u d Lake Ips&j Opax Mtn Q.  0.20  CO CD CL *  0.15  CD  £ CJ CD  0.10  o_  O) c  CO CD  0.05  0.00 Control  1 1  20%I.T.S.  20%P.C  50%I.T.S.  5 0 % P . C 50%I.T.S.(R)  Treatment  Figure 54: Mean specimen number (+/- s.e.) of C. unicolor for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=11.07; chi-square(mud)=4.05  (50%P.C.=50%I.T.S.(R)=Control=20%I.T.S.=20%P.C.=50%I.T.S.),  chi-square(opax)=1.97 (20%P.C.=50%I.T.S.=50%LT.S.(R)=50%P.C.=20%I.T.S.=Control).  0.40  0.35  co"  Mud Lake  1  Opax Mtn  0.30  03  0.25 H c  CD  E  0.20 H  o CD  S" 0.15-1 c  CO  I  0.10  I  X  0.05  0.00  Control  20%I.T.S.  20%P.C.  50%I.T.S.  5 0 % P . C . 50%I.T.S.(R)  Treatment  Figure 55: Mean specimen number (+/- s.e.) of S. americanus for seasons 1995-1997 (Mud Lake and Opax Mtn.), based on spring samples only. Chi-square(crit.)=l 1.07; chi-square(mud)=18.49 (20%I.T.S.=50%I.T.S.), chi-square(opax)=20.14 (20%I.T.S.=50%I.T.S.(R), 50%I.T.S.=Control).  106  DISCUSSION  The number of carabid beetle species collected at the Opax Mountain Silvicultural Systems Project research site was considerably greater than from comparable projects in various biogeoclimatic zones across B C (Craig 1995, Lemieux 1998, McDowell 1998, Lavallee 1999). A large number of traps, as well as a close proximity to a vagile grassland carabid fauna near Mud Lake, are two possible reasons accounting for the high species number in this study. Although 26 species found from this study were common to other studies (Craig 1995, Lemieux 1998, McDowell 1998, Lavallee 1999), 32 species were unique to the Opax Mountain Project research study. This study also recorded a geographic range extension for two species, namely Amara aenea, and Badister obtusus, both previously unrecorded from B C (Jarrett and Scudder 2001). Since this study, however, a re-identification of the carabid beetles from the Spencer Entomological Museum at U B C , has revealed that A. aenea was previously collected in B C back in 1951 (Jarrett and Scudder 2001). A carabid community composed of a high percentage of 'winged' species usually suggests a more disturbed or unstable habitat, where beetles invest heavily on flight (den Boer 1970). In this study, almost 85% of the carabid species are capable of flight, meaning they are either macropterous, dimorphic, or polymorphic with respect to wing morphology. Compared to other carabid beetle studies across B C (Craig 1995, Lemieux 1998, McDowell 1998, Lavallee 1999), the percentage of 'winged' species at the Opax Mountain Project research area is considerably higher. Because of the shorter natural disturbance return interval of IDF forests (B.C. Ministry of Forests and B.C. Ministry of Environment, Lands and Parks 1995), as well as the fact that some of the stands in these  107  forests have been logged three times in the past 90 years (Chan 1987, Vyse et al. 1991), a carabid assemblage composed of a high number of 'winged' species is not surprising. Assuming that control blocks represent pre-harvest carabid communities in spring samples, the majority of species arriving after logging occurred (immigrant species) dispersed primarily by flight (Tables 5 and 6). For example, only one (Calosoma wilkesii) of the 28 immigrant species from the Mud Lake site, and two (Scaphinotus relictus, and Pterostichus riparius) of the fourteen immigrant species from the Opax Mountain site, are known to be brachypterous (Lindroth 1961, 1963, 1969). Thus, while the possibility exists that carabids might use the extensive road system for dispersal into logged sites, this does not appear to be the case in this study, with most species probably arriving by flight. In addition, none of the dominant brachypterous carabid species found in this study (Table 1) were restricted to any of the six treatments, but were either capable of dispersing into the surrounding treatments on foot after logging, or survived in situ in the logged treatments. Even though there was a lack of buffer zones between treatments, corridors linking uncut habitats in control and patch-cut treatments still existed for those brachypterous species unwilling to traverse clearcuts (Figure 2). Also, three species of carabid (Loricera pilicornis pilicornis, Badister obtusus, and Trechus tenuiscapus), comprising four individuals, were found only in pre-harvest fall samples, and nowhere else (Table 1). One other species, Trichocellus cognatus, was also found exclusively in pre-harvest samples, but only at the Opax Mountain site (Table 3). This might simply reflect the fact that all five traps were sorted in pre-harvest fall samples, while only two out of the five were sorted in subsequent post-harvest fall  108  samples, leading to the possibility that these species might have occurred in the remaining unsorted post-harvest samples. Nevertheless, with only a few individuals of these four species, it is difficult, if not impossible to know whether the changes in the habitat due to the various logging practices was the main factor causing these species to disappear in post-harvest samples. Given the close proximity of the research area to the city of Kamloops, it was surprising to find four alien species, comprising a total of only five individual beetles. Whether alien species of carabids are out-competed in these forests, ill adapted to habitat, or have yet to extend their range, is still unclear.  M u d Lake and Opax Mountain Based on the total collection from the years 1993-97, the Opax Mountain site habitat was considerably more favorable, in terms of number of beetles collected, with a total of 26,185 beetles (Table 3) compared to only 16,734 from Mud Lake (Table 2). This was especially noticeable in the dominant carabid species numbers. This difference could reflect a situation where competition and predation of carabids is low at the Opax Mountain site, resources are plentiful, the preferred microclimate for the dominant species is more abundant, or conceivably some combination of these factors. In terms of numbers of species, the Mud Lake site was the more diverse with 49 species (Table 2), compared to only 36 from Opax Mountain (Table 3). This was surprising considering Mud Lake had almost 10,000 fewer beetles than Opax Mountain. The high number of grassland or open habitat species appearing at the Mud Lake site after logging, compared to Opax Mountain, is the main difference. In fact, of the 22 species unique to  109  Mud Lake, only one species is found exclusively in forests, with the rest known to inhabit meadows, grasslands, cultivated lands, tundra, riparian areas, or other open habitats. This was not the case at Opax Mountain, however, where most of the unique species are known only from riparian and forested habitats. Whether the Mud Lake site was more favorable to the grassland/open habitat species, or simply reflected an adjacent Bunchgrass ecosystem with a vagile grassland/open habitat carabid fauna, is still unclear. Thus, the Opax Mountain site is probably more representative of carabid diversity for LDFdk forests in southern B C , with Mud Lake assemblages being more of a mixture of forest and grassland species, and typical of the IDFxh biogeoclimatic zone. In response to the various harvesting treatments, the carabid assemblages at Mud Lake and Opax Mountain did show some disparities, although for the most part responded very similarly. For percent removal (20% vs. 50%), both sites generally recorded higher values for 50% removal treatments for all indices (spring and entire year samples) with the following exceptions: number of individuals (spring and entire year samples at both Mud Lake and Opax Mountain), Simpson's index (spring and entire year samples at Mud Lake, spring samples at Opax Mountain), Shannon-Wiener index (spring and entire year samples at Opax Mountain, entire year samples at Mud Lake), and species richness (spring and entire year samples at Opax Mountain) (Tables 7-10). As for the response to method of harvesting (I.T.S. vs. P . C ) , both sites showed patchcut treatments to yield higher numbers for all indices (spring and entire year samples), with the following exceptions: number of individuals (spring and entire year samples at both sites), number of species (spring and entire year samples at Mud Lake), and  110  Shannon-Wiener and Simpson's indices for spring samples at Opax Mountain (Tables 710). In response to leaving 30% of the treatment block in uncut reserves around snags, large veterans, and broadleaf groups, 50% I.T.S.(R) treatments recorded values that were for the most part intermediate between 20% and 50% I.T.S. treatments for all indices (spring and entire year samples), at both Mud Lake and Opax Mountain sites (Tables 7-10). Exceptions to this were the following: species number (spring and entire year samples at Opax Mountain, spring samples at Mud Lake), Shannon-Wiener index (entire year samples at both sites), species richness (spring and entire year samples at Opax Mountain) and Simpson's index values in entire year samples at Mud Lake. There are a number of possible reasons for the differences seen between the two sites in their response to the various harvesting treatments. One major difference is that they are situated in different biogeoclimatic subzones. Although Lemieux (1998) found biogeoclimatic subzones to correlate poorly with carabid fauna in northern BC's ESSF biogeoclimatic zone, previous research on carabids has shown that temperature (Mitchell 1963), soil moisture (Thiele 1977), and altitude (Eyre and Luff 1994), do in fact affect carabid distribution. Also, slight differences in vegetation, pest incidence, grazing and logging history might have caused the composition of carabid species in the two sites to respond differently to the various harvesting treatments. Because of these apparent differences, each block was analyzed separately in the rest of the study.  Ill  S p r i n g vs. entire year sampling  In comparing spring to entire year samples at both Mud Lake and Opax Mountain, all indices generally showed the same pattern (highest and lowest index values), in response to various harvesting treatments. The only major exceptions to this was for the Simpson's diversity index and number of species at Mud Lake (Tables 7-10). Although I chose to use spring samples to analyze in depth for this study based on their higher abundance and greater species number, the results from Tables 7-10 clearly show that spring and entire year samples respond for the most part similarly to the various harvested treatments, although with a slightly different fauna composition.  Species Richness  The results from Rarefaction shows that logging corresponded to an increase in species richness in all treatments during the three-year post-harvest period, compared to uncut control blocks (Figures 4-6). These results agree with previous work done on carabids, and their response to forestry practices (Lenski 1982b, Niemela and Halme 1992, Niemela et al. 1993a, Duchesne and McAlpine 1994, Spence et al. 1996, Lavallee 1999). This increase in species richness is believed to be the result of an increase in habitat heterogeneity (Southwood et al. 1979, Haila et al. 1994, Halme and Niemela 1993), a generally higher 'biological productivity' (Halme and Niemela 1993), as well as the disruption of competitive exclusion (Lenski 1982a & b), that occurs after logging. However, the method used to correct for different sample sizes, Rarefaction, was based on a sample size of only 5 for 1996, which might be too small to give reliable results  112  (Simberloff 1978). Nevertheless, the results do show a trend of increased species richness when uncut blocks were harvested. In examining each of the treatments for the three post-harvest years, the treatment showing greatest richness in each site changed slightly from year to year. At Mud Lake it was the 50% LT.S.(R) treatment for 1995, 50% patch-cut treatments in 1996, and then the 50% I.T.S.(R) again for 1997. At Opax Mountain, the 20% patch-cut treatment was highest in richness for 1995, followed by the 50% patch-cut treatment in 1996 and 1997. If richness increased owing to disturbance at these two sites, than Mud Lake was clearly the most disturbed site. Only in the 50% patch-cut treatment at Opax Mountain in 199697, was the disturbance level high enough to result in an increase in richness comparable to the more disturbed treatments at Mud Lake. This would suggest a more stable carabid community at the Opax Mountain block, than that seen at Mud Lake. Overall, Mud Lake and Opax Mountain display relatively similar values for richness in the first two post-harvest seasons of this study, with only a few minor exceptions. However in the 1997 season, the Mud Lake site responded more strongly to harvesting in all treatments except 20% I.T.S., with a substantial increase in richness compared to control blocks. Thus, while the affects of logging on carabids may have stabilized for the Opax Mountain carabid assemblage, there were continued changes in the Mud Lake assemblage, three years after harvesting. Looking at the temporal change in richness over the three-year period, a marked decrease in richness was observed for the 1996 season. One possible reason for this decrease in richness, was that trapping began two weeks earlier in 1996, owing to an early snowmelt. B y setting the traps two weeks earlier, it's possible that the carabids  113  were not as active as they might have been in the following two weeks, causing a decrease in the abundance, and hence richness, of the specimens collected in 1996.  Evenness  The species abundance distribution that most closely resembles the data at both sites for the post-harvest years 1995-97 (Figures 7-12) is the log-series. This pattern of species distribution would be predicted to occur in a situation in which species arrived in an unsaturated habitat at random intervals of time, to occupy similar fractions of the remaining niche hyperspace (Boswell and Patil 1971, May 1975). Evenness in the carabid community can be extracted from these graphs, by observing the slope of the underlying species abundance plot line. For example, a steep slope would indicate a community that is dominated by a few species with the remainder being fairly uncommon (the geometric series), while a rather flat sloped plot line (the broken-stick model) would indicate a situation where species are equally abundant. The log-series model observed from this study is similar to the geometric series model, differing only in an increase in the number of intermediate abundant species. The carabid community in spring 1995 showed relatively minor differences between treatments in both the Pielou's J's evenness measure, and the species abundance distribution plot lines. However, differences in community evenness start to become apparent in 1996, with both patch-cut treatments at Mud Lake, and 50% patch-cut treatments at Opax Mountain becoming more even. The Whittaker plot describes this more clearly by showing graphically that the intermediate abundant species are responsible for this increase in evenness. For 1997, little change occurred at the Opax  114  Mountain site from the previous year. The Whittaker plots for Mud Lake in 1997, however, do show changes in evenness from the previous year, as do the evenness measures of Pielou's J ' . The 50% I.T.S. treatments resulted in an increase in evenness at Mud Lake, which can be once again explained by the increase in the number of intermediate abundant species, as shown by the Whittaker plots. The 20% I.T.S. treatments, however, resulted in a decrease in evenness over the past year. Although a decrease in the slope of the 20% I.T.S. treatment plot line is not readily apparent, an increase in the number of rare species over the past year resulted in a lower evenness measure, as shown in the Pielou's J ' index. Also at Mud Lake in 1997, the Pielou's J ' index and Whittaker plots both show a decrease in evenness in patch-cut treatments from the previous year. Logging practices at both sites resulted in changes in the evenness of carabid communities, in much the same way as they did for richness measures. The increase in evenness, as depicted in the Whittaker plots for 1995-97, is once again more apparent in the Mud Lake site, with all but 20% I.T.S. treatments showing a greater evenness than control blocks. The Whittaker plots of Opax Mountain for the same time period, however, show that logging evidently only affected the 50% patch-cut treatments, with both intermediate and common species increasing in abundance. This same trend is apparent in the Pielou's J ' index, but not as strong. Reasons for the increase in evenness could possibly come from a decrease in the amount of competition for resources (Southwood 1978, Magurran 1988), and/or changes in microclimate resulting from logging (Thiele 1977). Until experimental work is done to elucidate the reasons for this,  115  one can only speculate as to what's happening in carabid communities, when forests are logged.  Heterogeneity  The heterogeneity of carabids was investigated using two non-parametric measures of diversity, namely the Shannon-Wiener and Simpson's diversity index, as well as the logseries based alpha diversity index. With only a few exceptions, owing to the different weighting schemes of each index, all three heterogeneity indices responded similarly to the changes in the carabid assemblages resulting from the various harvesting practices. The general trend of carabid species diversity increasing as a result of forest harvesting (Bultman et al. 1982, Lenski 1982b, Boyle 1991, Niemela et al. 1992, Niemela et al. 1993b, Craig 1995, McDowell 1998, Lavallee 1999) was also observed in this study. The only significant differences in heterogeneity, as measured by the Shannon-Wiener and Simpson's index in 1995 (Figures 19 and 20), was the significantly greater heterogeneity of 20% patch-cuts compared to 50% I.T.S.(R) treatments at the Opax Mountain site. Although the alpha index showed a greater difference between 20% P.C. and 50% I.T.S. treatments at Opax Mountain (Figure 16), large standard errors negated any statistical significance. Of the three indices, the alpha index showed the greatest difference between treatments at Mud Lake in 1995, with an obvious trend of greater heterogeneity, though not statistically significant, for the higher percent removal 50% I.T.S., 50% P . C , and 50% I.T.S.(R) treatments. This differed from Shannon-Wiener and Simpson's indices, which had 50% I.T.S. and 50% P.C. treatments not as diverse, but otherwise showing the same pattern of response to logging. Control blocks at both Mud Lake and Opax Mountain contained the least diverse carabid assemblage overall, with  116  only 50% I.T.S.(R) (Shannon-Wiener and Simpson's index), and 50% I.T.S. treatments (alpha index) at Opax Mountain with a slightly lower heterogeneity measure. Except for the unusually low value of heterogeneity recorded by the Shannon-Wiener index for the 50% I.T.S. treatment at Mud Lake in 1996, all three indices showed a very similar response to logging, with an increase in heterogeneity in the 50% I.T.S., 20% and 50% patch-cut treatments (Figures 17, 21 and 22). The only significant difference detected at Mud Lake for 1996, was the significantly greater heterogeneity of 50% patchcuts compared to 50% I.T.S. treatments, as measured by the Shannon-Wiener index. Because the Shannon-Wiener index is more sensitive to changes in the rare species, a reduction in evenness and/or richness of the rare carabid species must have occurred in the 50% I.T.S. treatment. For the common carabid species, however, increases in evenness and/or richness were responsible for the high heterogeneity values recorded in 50% I.T.S. and patch-cut removal treatments. In addition to this, both Shannon-Wiener and Simpson's indices recorded a marked drop in heterogeneity, from the previous year, for 20% I.T.S. and 50% I.T.S.(R) treatments at Mud Lake. At Opax Mountain in 1996, smaller standard errors in each of the treatments resulted in A N O V A tests detecting more significant differences between treatments for each of the three indices, though to a greater extent in the Shannon-Wiener and Simpson's, than in the alpha index. Thus, indices for the 50% patch-cuts were significantly greater than control blocks (all three indices), 20% patch-cuts (Shannon-Wiener and Simpson's indices), and 50% I.T.S.(R) treatments (Shannon-Wiener index), while 20% I.T.S. treatments were significantly greater than control and 20% patch-cut treatments (Shannon-Wiener and Simpson's indices). The same pattern of response to logging was observed in each of the three  117  indices, with an increase in diversity for 20% I.T.S. and 50% patch-cut treatments, and a substantial decrease in diversity for 20% P.C. and 50% I.T.S. treatments. Control blocks recorded the lowest values of heterogeneity in 1996, with only the 50% I.T.S. treatment (Shannon-Wiener index) and 20% I.T.S. treatment (Simpson's index) at Mud Lake recording a slightly lower value. Because of the earlier trapping period in 1996, the Shannon-Wiener diversity index measured a drop in diversity at both the Mud Lake and Opax Mountain site. In 1997, the only major change in carabid diversity at Mud Lake occurred in the 50% I.T.S.(R) treatment, which increased substantially in all of the three indices from the previous year (Figures 18, 23, and 24). This resulted in 50% LT.S.(R) treatments having a significantly greater heterogeneity than control blocks (Shannon-Wiener and alpha indices), and 20% I.T.S. treatments (Shannon-Wiener index), while 50% patch-cut treatments were significantly greater than control and 20% I.T.S. treatments (ShannonWiener index). At Opax Mountain in 1997, diversity indices in the 50% patch-cut treatments increased, while 20% I.T.S. treatments slightly decreased in diversity over the previous years values, with other treatments remaining near 1996 diversity levels. Because of the small decrease in diversity observed for 20% I.T.S. treatments at Opax Mountain, fewer statistically significant results were detected between treatments, compared to 1996. Control blocks at Mud Lake are once again the least diverse of the six treatments, with only 20% I.T.S. treatments slightly lower for the Simpson's index measure of diversity. At Opax Mountain, all three indices showed a marginal increase in diversity in the control block over the previous year's value. Control blocks now were 3  118  rd  lowest in Shannon-Wiener and Simpson's indices, behind 20% P.C. and 50% I.T.S., and second lowest behind only 50% I.T.S. for the alpha statistic measure of diversity. In comparing Mud Lake to Opax Mountain, logging evidently had more of an impact on carabid heterogeneity at the lower elevation Mud Lake site, though this is not readily apparent until the 1996-97 season. Similar to other studies (Lemieux 1998, McDowell 1998, Lavallee 1999) logging resulted in changes in the more common species, as measured by the Simpson's index, to be greater than those for the less common species, as measured by the Shannon-Wiener index. Patch-cut methods resulted in higher heterogeneity measures than individual-tree selection methods. However, this is mostly apparent in 1996-97 for Mud Lake blocks, with little or no trend seen at Opax Mountain. Other than the low value in 50% I.T.S. treatments at Mud Lake in 1996 (Shannon-Wiener index), 50% removal treatments resulted in a greater heterogeneity for carabid assemblages in the 1996-97 season. As with method of harvesting (I.T.S., I.T.S.(R), and patch-cuts), percent removal (20% and 50%) differences are showing no obvious trend at the Opax Mountain site. Thus, the carabid assemblages at Mud Lake appear to have been impacted more heavily by logging, than those at Opax Mountain, and this was especially noticeable from 1996 on.  T a x o n o m i c distinctness  Taxonomic distinctness indices differ from standard species diversity measures in that they factor in the phylogenetic structure of each species, or individual, and compare this with others in the same sample, measuring the 'distance' between them. Thus, not only can biological communities be measured for their diversity of species, but also for their  119  diversity of evolutionary relationships in the individuals and species. One advantage of using taxonomic distinctness measures is that they can differentiate between communities that share a similar species diversity, by describing the 'relatedness' of the components of that diversity. One other benefit of using distinctness measures is that they are not influenced by small sample sizes, which can pose problems for regular diversity measures (Warwick and Clarke 1995). In this study, distinctness was measured from a master list of species collected in the Opax Mountain Project area, and branch lengths were measured using species, genera, tribe, and subfamily taxonomic groupings. In addition to the distinctness measures, 'probability funnels' were produced which allowed for comparisons of sample distinctness with 'expected' distinctness values, taken from the master list of species at the Opax Mountain Project research study. Thus, not only were samples tested for differences between treatments, but also for differences between treatments and 'expected' values. The results for taxonomic distinctness (TD) in 1995 (Figure 25) show control, 50% patch-cut, and 20% I.T.S. treatments significantly greater than 50% I.T.S. treatments at the Opax Mountain site. This contrasts with average taxonomic distinctness (aveTD) (Figure 28), which shows 50% patch-cut treatments significantly greater than 50% I.T.S. treatments at the Mud Lake site. The results from the variation in taxonomic distinctness (varTD) (Figure 31) shows the opposite result to aveTD, with 50% I.T.S. treatment values significantly greater than 50% patch-cuts at the Mud Lake site. T D indices also record the highest distinctness values for control treatments, where aveTD and varTD indices show control treatments as one of the least distinct communities.  120  Results in 1996 for TD (Figure 26) show that control, 50% patch-cut, and 50% I.T.S.(R) treatment values are significantly greater than 50% I.T.S. treatments, for the Opax Mountain site. No significant differences were detected for aveTD in 1996 (Figure 29). However, values for 20% patch-cuts were significantly greater than control blocks at the Mud Lake site, for varTD (Figure 32). Compared to the previous year at Mud Lake, both TD and aveTD dropped slightly in control and 20% I.T.S. treatments. However a more substantial decrease, detected by all three distinctness indices, occurred in 50% I.T.S. treatments, and in control blocks for varTD at Mud Lake. A drop in the number of rare grassland genera might have caused of this decline, as the ShannonWiener index, which is sensitive to changes in the rare species in the community, also shows heterogeneity to drop off for this same time period. At Mud Lake in 1997, all three distinctness indices show treatment values returning to their 1995 levels, with only minor exceptions (Figures 27, 30, and 33). Results at Opax Mountain for 1997 show T D and aveTD relatively unchanged from the previous year. Although a statistical significance was detected for varTD in 1997 at Opax Mountain, the Tukey test was unable to determine which sites were significantly different from others. Over the three-year study, carabid assemblages at Opax Mountain were generally more taxonomically distinct than their Mud Lake counterparts, for both TD and aveTD. For varTD, the opposite trend was apparent, with Mud Lake recording greater values for most of the treatments in the three-year study. One possible reason for this difference is that most of the new species at Mud Lake came from only two genera, Amara Bonelli and Harpalus Latreille, which might increase species diversity, but not necessarily distinctness.  121  Assuming a null hypothesis that each treatment contains species randomly selected from the total species list, neither aveTD nor varTD values for both Mud Lake and Opax Mountain carabid assemblages (Figures 34-36), fell outside the 95% confidence funnel, using presence/absence data. Taxonomic distinctness indices were introduced in 1995 by Warwick and Clarke as a more sensitive univariate index of community perturbation than standard species diversity. Similar to marine communities under perturbed situations (Warwick and Clarke 1995), the epigeal carabid communities, at both Mud Lake and Opax Mountain (Figure 25 &27), showed a decrease in T D when forests were logged. Results from T D indices for 1996 (Figure 26), as well as from both aveTD and varTD indices (Figures 2833), however, shows no such trend. Because this study represents the first of its kind in using taxonomic distinctness indices to measure changes in a perturbed carabid beetle community, further work on using carabids from different ecosystems would be beneficial in understanding more thoroughly, the changes in carabid assemblages that may result from logging practices.  M u l t i v a r i a t e analysis  The most obvious pattern observed from this study, for both the M D S and cluster analysis, is the clear demarcation between Mud Lake and Opax Mountain carabid assemblages. Only in control and 20% LT.S. treatments for 1995 (Figures 40 and 41), and in control blocks for 1997 (Figures 44 and 45) at Mud Lake, do carabid assemblages resemble those from Opax Mountain. These results suggest that different biogeoclimatic subzones exert a stronger influence on the composition of carabid assemblages, than do  122  treatment differences. Whether this is owing to differences in vegetation, food supply, microclimate, pest incidence, grazing and logging history, the closer proximity of Mud Lake to the Bunchgrass biogeoclimatic zone, or from simply stochastic reasons, is still unclear. Another pattern observed from both M D S and cluster analysis, is the higher percentage similarity and closer proximity, between the various harvesting practices at Opax Mountain, than for those at Mud Lake. Similar to the univariate results mentioned earlier, the impact of logging on carabid beetle assemblages appear to have influenced the Mud Lake assemblage to a greater degree, resulting in more distinct differences between treatments, than between those seen at Opax Mountain. With the odd exception, the method of harvesting generally appears to have exerted a greater influence over carabid assemblages, than did percent removal, with both patch-cut and I.T.S. treatments clustering and mapping closer together, than to treatments with a similar percentage removal. Thus, carabid assemblages appear to be responding more to forest type, or successional stage, than to similar amounts of crown removal, using a different method of harvesting. This is probably because of the occurrence of different carabid feeding guilds within the two methods of harvest, with generalist ground predators in I.T.S. treatments, and seed feeding herbivores (at least in the adult stage) occupying patch-cut treatments. Both M D S and cluster analysis also show carabid assemblages from control blocks for the most part resembling I.T.S. over patch-cut methods of forest removal. These results agree with what McDowell (1998) found in both ESSF and ICH biogeoclimatic zones, with old growth carabid assemblages clustering closer to partially cut sites than to clearcut sites.  123  Dominance structure The dominance structure at Mud Lake (Tables 11 &12) showed that spring and entire year samples shared seven of the top ten most abundant carabid species found in control blocks. The top four species from spring samples were also the top four species from entire year samples, but switched only in dominance position and percent abundance. The three species that were found in higher positions of dominance in spring samples but disappeared entirely from the top ten list in entire year samples, were all spring breeders, while two of the species that replaced them were autumn breeders. Interestingly, Synuchus impunctatus, the sixth most abundant species from entire year samples, was totally absent from spring samples. In contrast to this, all of the top ten species from spring samples at Opax Mountain were on the top ten list for entire year samples (Tables 13&14). The dominance structure that most closely resembled that observed from control blocks was 20% I.T.S. treatments for both spring and entire year samples at Mud Lake. Although logging impacted the carabid fauna to a lesser extent at Opax Mountain than at Mud Lake, 20% I.T.S. treatments were also the closest in resemblance to control block dominance structure for entire year samples, while 20% patch-cut treatments resembled control block structure for spring samples. Thus for the most part, carabid assemblages from lower percent removal treatments, as well as from individual-tree selection methods, showed the most resemblance to the dominance structure from control blocks. Both sites also showed that the most dominant 4-5 species from control blocks, remained the most dominant 4-5 species in all harvested treatments, while most of the  124  changes in species positioning and turnover happened in the bottom 5-6 species. Although the number of individuals in the dominant 4-5 species sometimes changed substantially in the different harvested treatments, it was because of their relatively higher abundance that they appeared to be more stable, while the less abundant species from the top ten list, which often only changed slightly in abundance in the various harvested treatments, appeared relatively less stable in dominance position.  Species analysis The three habitat-types, created in the patch-cut and 50% I.T.S.(R) treatments (uncut, edge, and clearcut), were combined together in this study into a single treatment-type. Although this prevented me from grouping species into their various habitat-type preferences, as found in other studies (Niemela et al. 1992, Halme and Niemela 1993, Craig 1995, McDowell 1998, Lemieux 1998), I was still able to group the ten most abundant species (>200) into their respective removal level and method of harvest preferences. The groupings were as follows; 1) uncut forest species, 2) medium percent removal (with reserves) species, 3) high percent removal species, 4) patch-cut species, 5) individual-tree selection species, 6) generalist species and, 7) unresolved species. Species were placed into the various groupings depending on whether A N O V A tests showed any significance, or if any recognizable trend in the species preference was observed. The reasoning behind combining habitat-types into a single treatment-type was because of the small size of clearcuts and reserves, which could be easily traversed by most carabid species (Thiele 1977), as well as insufficient information on the size of clearcuts and reserves the traps were placed in. A further reason for grouping the habitat-  125  types into a single treatment-type, was that a separate "edge effects" study was set up to examine the response of ground beetles to edge habitats. Uncut forest species Two species standout as preferring uncut forests to other logged treatments. The first is Carabus taedatus agassii, which although abundant in every treatment, showed the highest mean number of individuals per trap in uncut control blocks at both sites, though not statistically significant at the Mud Lake site (Figure 47). Lindroth (1961) describes C. taedatus agassii as a xerophilous species, preferring open gravelly soil with low vegetation, and occurring in open coniferous forests. Frank (1971) has also recorded C. taedatus agassii from an arable field habitat. However, Craig (1995) found the subspecies C. taedatus taedatus to be a potential old growth species in the C W H forests on Vancouver Island, but was unsure of this association owing to poor replication. I have also captured C. taedatus agassii individuals in the Bunchgrass biogeoclimatic zone in the southern interior of B C , a habitat that differs considerably from Craig's old growth forest classification. Data from this study supports Craig's data to a certain degree, in that uncut forests contain more individuals. However, because species in the genus Carabus Linnaeus are proficient walkers (Lindroth 1961) capable of dispersing great distances (especially in open LDF forests), a high number of beetles might have dispersed from uncut areas into the surrounding treatments. Thus, while it appears that the sub-species C. taedatus taedatus prefers the old growth forests along the coastal forests of B C (Craig 1995), the sub-species C. taedatus agassii found in this study, is more xerophilous as noted by Lindroth.  126  The other carabid species to significantly occur more often in uncut over logged forests was Calathus ingratus at Mud Lake (Figure 49). Lindroth (1966) found this species to prefer moderately moist or rather dry open gravelly ground in deciduous forests. Other studies describe C. ingratus as a generalist species, occurring in equal numbers in a variety of forest types and edges (Niemela et al 1993a, Niemela 1997). Although dimorphic in wing morphology (Lindroth 1966), dispersal is probably carried out by brachypterous specimens, as Lindroth found the macropterous morphs to be rare.  High removal species Pterostichus adstrictus at Opax Mountain was the only dominant carabid species captured most abundantly in the more disturbed higher removal treatments (Figure 50). Liemeiux (1998) found P. adstrictus species to occur mostly in disturbed habitats in northern ESSF zones, calling it a disturbance specialist. McDowell (1998) also found P. adstrictus mostly in disturbed clearcut habitats, in both the ESSF and ICH forests in B C . Research into the effects of logging practices in Alberta, has also shown P. adstrictus to occur more frequently in early successional forests to primary forests (Niemela et al 1993a & b).  Medium removal (with reserves) species Three carabid species in this study were found to occur predominantly in the medium removal (with reserves) treatments. These three species were Pterostichus adstrictus at Mud Lake, and Scaphinotus marginatus and Calathus advena at Opax Mountain. For Pterostichus adstrictus, the 50% LT.S.(R) treatment contained the greatest mean number of specimens per trap, with 50% patch-cut treatments slightly less abundant  127  (Figure 50). Although adults of P. adstrictus have previously described as a disturbance species in this and other studies (Niemela et al. 1993a & b, Lemieux 1998, McDowell 1998), P. adstrictus larvae have been found to rely on the microclimate provided by coarse woody debris (Goulet 1974). Thus, the high number of specimens caught in 50% I.T.S.(R) treatments could possibly stem from the fact that large logs were left around old-growth trees, providing a critical habitat for P. adstrictus larvae. Scaphinotus marginatus populations at Opax Mountain also occurred most often in medium percent removal (with reserves) treatments compared to other treatments (Figure 51). Lindroth (1961) describes S. marginatus as a rather eurytopic species, occurring in forest habitats in the southern latitudes of Canada. Research in the forests of Alberta has shown S. marginatus to be a forest generalist (Niemela et al. 1992, Spence et al. 1996), and an old growth coniferous specialist (Niemela et al. 1993a & b, Spence et al. 1996), with no urge to cross edge habitats into clearcuts (Niemela 1997). In B C studies, Craig (1995) was unsure as to its habitat preference, Lemieux (1998) classified 5. marginatus as a generalist species, while McDowell (1998) called it a forest species. Thus, Lindroth's description of S. marginatus von as a eurytopic species seems to be a very apt one. The final species to occur most abundantly in medium removal (with reserves) treatments compared to others was Calathus advena at Opax Mountain (Figure 48). This ground beetle has been described in other studies as a forest species (Lindroth 1966), an old growth species (Niemela et al. 1993b, Spence et al. 1996) a generalist species (Lemieux 1998), and a clearcut species (Duchesne and McAlpine 1992). As with Scaphinotus marginatus, relative abundance of Calathus advena appears to result from  128  some interaction between the species and characteristics of the forest type itself (Spence et al. 1996), rather than a specialization to any particular habitat-type in itself.  Patch-cut species Patch-cut species are those species that are generally regarded in other studies as open habitat species, overwintering in the adult stage, active in sunny daylight hours, and proficient fliers (Thiele 1977). Three species of ground beetles from this study were recognized as patch-cut species, namely Bembidion dyschirinum, Notiophilus directus, and Syntomus americanus. For Bembidion dyschirinum, a statistically significant higher number of specimens were captured in patch-cut treatments compared to other treatments, at both the Mud Lake and Opax Mountain site (Figure 52). Lindroth (1963) described this species as an open country species, which agrees with the findings from this study. Lindroth also believed the macropterous forms of this dimorphic species to be rare, but whether this was the case in this study is unclear, as wing morphology was not examined. Notiophilus directus at Opax Mountain was also found to occur more often in patch-cut treatments, though not significantly (Figure 53). Lindroth (1961) describes N. directus as a xerophilous and heliophilous species, but also mentions that the genera Notiophilus Dumeril often gives the impression that their habitat choice is due to mere chance. With N. directus at Opax Mountain in this study, this does not appear to be the case, as a greater number of beetles would be expected to occur in the open habitats created by patch-cutting, assuming Lindroth's habitat description is correct. McDowell (1998) also  129  found this species to occur mostly in clearcuts in her study in the ESSF and ICH forests of central B C . The third species found to significantly occur more often in patch-cut treatments at both the Mud Lake and Opax Mountain study sites was Syntomus americanus (Figure 55). Lindroth (1969) describes S. americanus as a sun loving, xerophilous species, found on sandy, rarely peaty soil, with sparse low vegetation. Craig (1995) found S. americanus to be a regeneration specialist from her study on Vancouver Island. I have also seen S. americanus in the grasslands just north of Calgary, Alberta, running alongside ants in the bright sunshine.  Individual-tree selection species The only dominant carabid species to be captured mostly in individual-tree selection treatments was Calathus ingratus at Opax Mountain (Figure 49). A statistical significance was detected only for the 20% I.T.S. treatment, with 50% I.T.S. treatments only marginally greater than 50% patch-cut treatments. An unexpected result for the Opax Mountain site was the low number of C. ingratus individuals collected in control treatments, where at Mud Lake the same species is described as a uncut forest species. Reasons for the disparity in control block numbers is still unclear.  Generalist species The carabid species Cymindis unicolor showed no significant differences between any of the treatments at either block, nor showed any recognizable pattern of percent removal or method of harvest preference (Figure 54). Lindroth (1969) describes this species as  130  xerophilous, preferring tree-less country and tundra habitats. Although no significant differences were detected, there was a suggested preference for logged treatments, over uncut control blocks at the Opax Mountain site. This result at Opax Mountain fits Lindroth's description for the species, however, at Mud Lake, only the 50% patch-cut treatments contain a greater abundance of C. unicolor individuals when compared to control blocks, with other treatments at equal or lower abundance's. One possible reason for the difference in responses of C. unicolor between the two sites, might be the potentially greater competition between C. unicolor and the more diverse and abundant open habitat species at Mud Lake, resulting in a lower overall abundance of C. unicolor at the Mud Lake site.  Unresolved species The four species that were unresolved with respect to possible preferences were the Mud Lake and Opax Mountain populations of Pterostichus neobrunneus Lindroth, and the Mud Lake populations of Calathus advena, Scaphinotus marginatus, and Notiophilus directus. Although some of these species did show treatment-type preferences, none showed any preference for either percent removal or method of harvest. The most abundant species overall in spring samples, Pterostichus neobrunneus, showed a statistically significant difference between every treatment at both Mud Lake and Opax Mountain sites (Figure 46). Not only did it not respond to the various logging treatments in a way that was consistent with the above groupings, but the responses of P. neobrunneus at the Mud Lake and Opax Mountain, were quite different. Lindroth (1966) describes P. neobrunneus as a woodland species, occurring on rather dry ground. McDowell (1998) placed P. neobrunneus in her habitat generalist category, while Craig  131  (1995), like myself, was unsure as to its habitat preference. Whatever the grouping, P. neobrunneus is one of, if not the dominant carabid beetle found in all six treatments, for both Mud Lake and Opax Mountain habitats. Both Scaphinotus marginatus and Calathus advena populations at Mud Lake (Figures 51 and 48) also showed statistically significant differences between treatments, and like Pterostichus neobrunneus, no trend in percent removal or method of harvest was observed. Although Notiophilus directus at Mud Lake was also placed in the unresolved species group (Figure 53), this was mainly because of the small number of beetles collected from the site.  Other species Although this study analyzed only the ten most abundant species found in spring samples, a few of the less abundant species also displayed some interesting results. For example Scaphinotus angusticollis, a species common in west coast forests, responded to logging practices in this study in a manner similar to other studies (Craig 1995, McDowell 1998, Lavallee 1999), with a greater proportion of the individuals occurring in uncut or lower percent removal individual-tree selection treatments. Three of the other less abundant species, Bembidion grapii, Amara ellipsis, and Amara laevipennis, displayed an obvious patch-cut treatment preference, while 20 of the 21 specimens of Carabus serratus were found exclusively in 50% I.T.S.(R) treatments. Because every method of harvest evidently benefited some species at the expense of others, no one treatment appears sufficient for the whole carabid community. Instead, a mix of harvesting methods that maintained the greatest number of all native forest  132  species, including sensitive species, and over an indefinite period of time, would seem to be the best strategy for the preservation of carabid biodiversity. Although this study does provide some useful information to forest managers, there's no substitute to in-depth autoecological studies.  133  1) While species richness, evenness, and heterogeneity values did increase for the most part in logged treatments, the predicted trend of diversity values increasing with an increasing percentage of forest removal, was observed (with the odd exception) only at Mud Lake in the 1996/1997 season for the three heterogeneity measures. As for distinctness values, only in TD indices did I observe (with the odd exception) the predicted decrease in distinctness as a result of logging. No such trend was observed, in any of the three distinctness measures, however, that showed distinctness values decreasing with an increase in percent forest removal. Thus, with the exception of heterogeneity measures, the results from this study do not support hypothesis 1. 2) As described above, control blocks for the most part recorded the lowest values for species richness, evenness, and heterogeneity. However, only in heterogeneity values for Mud Lake in the 1996/1997 season, did I observe the predicted trend of increasing diversity values as you move from control to individual-tree selection to patch-cut methods of harvesting forests. As mentioned above, control blocks recorded higher TD values (with the odd exception) than any logged treatment. However, no trend was observed that showed any of the three distinctness indices to decrease as you move from control to individual-tree selection to patch-cut methods of harvesting forests. Although the results from heterogeneity values support hypothesis 2, species richness, evenness, and distinctness values, for the most part, do not.  134  3)  For the most part, carabid assemblages within each site show patch-cut treatments to cluster/map together, I.T.S. carabid assemblages to cluster/map together, and control blocks to cluster/map closer to I.T.S. than patch-cut treatments. The results from multivariate analysis, therefore, support hypothesis 3. Results also showed for the most part a clear demarcation between Opax Mountain and M u d Lake carabid assemblages, as well as a greater similarity in carabid assemblages between the different treatment blocks at Opax Mountain, than at M u d Lake.  4) Dominance structure tables generally showed carabid assemblages from lower percent removal treatments, as well as those from individual-tree selection methods, to most resemble the dominance structure found in control blocks. These results, thus, lend support to hypothesis 4. 5) Few species specialized in any one treatment, instead, most species were found to occur i n all treatments. This was probably due to the high percentage of 'winged' species, as well as the extensive history of past disturbances i n this LDF biogeoclimatic zone. However, one species was captured i n greater numbers (P. adstrictus (Opax Mtn.)) and others in fewer numbers (C. taedatus agassii, C. ingratus ( M u d Lake)) in higher percentage forest removal treatments, one species was caught in relatively equal numbers across all treatments (C. unicolor), while others were found to be caught in greater numbers in patch-cut treatments (B. dyschirinum, N. directus (Opax Mtn.), and S. americanus). The results from intraspecific analysis, therefore, support hypothesis 5. Surprisingly, some species were also caught in greater numbers in I.T.S. (C. ingratus (Opax Mtn.)) and  135  I.T.S.(R) (P. adstrictus (Mud Lake), S. marginatus (Opax Mtn.), and C. advena (Opax Mtn.)) treatments.  Although the Opax Mountain Silvicultural Systems Project site was setup to primarily test the effects of harvesting on animal populations, there were a number of problems with this setup. For example, replicates were situated in different biogeoclimatic subzones, there was insufficient replication within similar biogeoclimatic subzones, there was a lack of buffer zones, and practically no pre-harvest collection data. However, some important research did occur, for example, carabid responses to forest harvesting were studied for the first time in LDF forests, two new carabid species for B C were discovered, and a good inventory was constructed of the carabid fauna for the region. 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J., U.S.A.  150  Appendix 1: Pitfall trap collection dates for Opax Mountain Silvicultural Systems Project  Series #  Traps set  Traps collected  0000 1000 2000 3000 4000 5000  29.Sept.93 15.Oct.93 3.Aug.94 18.Aug.94 early.Jan.95 9.May.95 (Mud Lake) 23.May.95 (Opax Mtn.) 24.May.95 (Mud Lake) 5.June.95 (Opax Mtn.) 8.Aug.95 22.Aug.95 2,Oct.95 16.Oct.95 18.Jan.96 22.April.96 (Mud Lake) 7.May.96 (Opax Mtn.) 7.May.96 (Mud Lake) 21 .May.96 (Opax Mtn.) 6.Aug.96 20.Aug.96 30.Sept.96 15.Oct.96 6.Jan.97 6.May.97 (Mud Lake) 20.May.97 (Opax Mtn.) 20.May.97 (Mud Lake) 3.June.97 (Opax Mtn.)  13.Oct.93 29.Oct.93 17.Aug.94 1.Sept.94 early.March.95 23.May.95 6.June.95 7.June.95 19.June.95 22.Aug.95 5.Sept.95 16.Oct.95 30.Oct.95 18.March.96 6.May.96 21. May.96 21.May.96 4.June.96 20.Aug.96 3.Sept.96 14.Oct.96 29.Oct.96 6.March.97 20.May.97 3.June.97 3.June.97 17.June.97  6000 7000 8000 9000 10000 11000 12000 13000 14000 15000 16000 17000 18000 19000 20000  151  

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