'Old-Growth' and Managed Forests in Southeastern Ontario

BIODIVERSITY OF SAPROXYLIC COLEOPTERA IN 'OLD-GROWTH' AND MANAGED FORESTS IN SOUTHEASTERN ONTARIO

Rebecca Zeran Department ofNatural Resource Sciences McGill University, Montreal September 2004

A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements of the degree of Master of Science

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LIST OF TABLES ...... v LIST OF FIGURES ...... vi LIST OF APPENDICES ...... vii ACKNOWLEDGEMENTS ...... ix PREFACE ...... x CONTRIBUTIONS OF AUTHORS ...... xi ABSTRACT ...... xii RÉSUMÉ ...... xiii

CHAPTER 1: Introduction and Literature Review General introduction ...... 1 Eastern Ontario forests ...... 2 Old-growth forests ...... 3 Downed woody debris ...... 5 Saproxylic beetles in forest management ...... 8 Fungus-insect relationships ...... 12 OBJECTIVES ...... 14 REFERENCES ...... 15

CONNECTING STATEMENT ...... 21

CHAPTER 2: Biodiversity of fungivorous beetles (Coleoptera) in managed and 'old growth' hemlock-hardwood forests in southeastern Ontario ABSTRACT ...... 22 INTRODUCTION ...... 23 MATERIALS AND METHODS ...... 25 Study Sites ...... 25 Sample Collection ...... 25 Coleoptera Identification ...... 26

11 Fugus and Coarse Woody Debris Inventory ...... 27 Data Analyses ...... 28 RESULTS ...... 29 DISCUSSION ...... 31 Comparison of Forest Types ...... 31 Species Diversity of Leiodinae ...... 33 Species Diversity of Other Fungivores ...... 34 Trapping Differences ...... 35 Conclusions: Fungus-insect Re1ationships and Forest Management...... 36 REFERENCES ...... 39

CONNECTING STATEMENT ...... 80

CHAPTER 3: Sap beetles (Coleoptera: Nitidulidae) in managed and 'old-growth' forests in southeastern Ontario, Canada ABSTRACT ...... 81 INTRODUCTION ...... 82 MATERIALS AND METHODS ...... 84 Study Sites ...... 84 -Sample Collection ...... 84 Nitidulid Identification ...... 85 Data Analyses ...... 86 RESULTS ...... 87 Richness, Abundance and Phenology ...... , ...... 87 Site Comparisons ...... 88 Species Composition ...... 88 Effects of Glischrochilus quadrisignatus ...... , ...... 89 DISCUSSION ...... 89 Richness, Abundance and Phenology ofNitidulidae ...... 89 Comparison of Forest Types ...... 91 Resolving Patterns: Collecting Methods and Species Identification ...... 92

III Conclusion: Forest Management and Nitidulidae ...... 94 REFERENCES ...... 95

CHAPTER 4: General Conclusion ...... 12 3

IV LIST OF TABLES

Table 2.1. Location, tree species, size, age and general description of study sites ...... 45 Table 2.2. Fungivorous Coleoptera collected at study sites, 2003 ...... 46 Table 2.3. Raw species richness (Sobs), number ofindividuals, rarefaction estimated species richness (species ± sn, standardized to 1550 individuals), and diversity indices of fungivorous Coleoptera at study sites (both trap types pooled) ...... 47 Table 2.4. Indicator species analysis of fungivorous Coleoptera collected in flight- intercept traps ...... 49 Table 3.1. Location, tree species, size, age and general description of study sites ...... 100 Table 3.2. Raw species richness (Sobs), number ofindividuals, rarefaction estimated species richness (species ± sn, standardized to 120 individuals), and diversity indices of sap beetles (Coleoptera: Nitidulidae) at study sites (both trap types pooled) ...... 101 Table 3.3. Indicator species analysis of sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps ...... 102 Table 3.4. Sap beetles (Coleoptera: Nitidulidae) collected at study sites, 2003 ...... 103

v LIST OF FIGURES

Figure 2.1. Location ofstudy sites, eastem Ontario, Canada ...... 50 Figure 2.2. A - Flight-intercept trap (FIT); B - Trunk-window trap (TT) ...... 51 Figure2.3a. Rarefaction estimates of expected species richness (± 1 SD) of fungivorous Coleoptera collected with flight-intercept and trunk-window traps in six forest sites ...... 52 Figure 2.3b. Rarefaction estimates of expected species richness (± 1 SD) of fungivorous Coleoptera collected with flight-intercept and trunk-window traps in managed versus 'old-growth' forests ...... 53 Figure 2.4. Dendrogram of c1uster analysis on species of fungivorous Coleoptera collected from flight-intercept traps ...... 54 Figure 3.1. Location of study sites, eastem Ontario, Canada ...... 104 Figure 3.2. Phenology of selected Nitidulidae collected with both flight-intercept and trunk-window traps. A - Glischrochilus quadrisignatus (Say); B - Glischrochilus sanguinolentus (Olivier) ...... 105 Figure 3.3. Phenology of selected Nitidulidae collected with both flight-intercept and trunk-window traps. A - Cychramus adustus Erichson and Pal/odes pallidus (Palisot de Beauvois); B - Omosita colon (L.) ...... 106 Figure 3.4. Phenology of the most abundant species of Epuraea collected with both flight-intercept and trunk-window traps. A - Epuraea ru/a and E. labilis; B - E. erichsoni, E. depressa, and E. rufida ...... 107 Figure 3.5a. Rarefaction estimates of expected species richness (± 1 SD) of sap beetles (Coleoptera: Nitidulidae) collected with flight-intercept and trunk-window traps in six forest sites ...... 108 Figure 3.5b. Rarefaction estimates of expected species richness (± 1 SD) of sap beetles (Coleoptera: Nitidulidae) collected with flight-intercept and trunk-window traps in managed versus 'old-growth' forests ...... 109 Figure 3.6. Dendrogram of c1uster analysis on species of sap beetles (Coleoptera: Nitidulidae) collected from flight-intercept traps ...... 110

vi LIST OF APPENDICES Appendix 2.1. Fungi found within an approximate 25m radius of the flight-intercept traps at each site ...... 55 Appendix 2.2a. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in 'old-growth' forest #1, Cornwall, ON, (OG-l) ...... 56 Appendix 2.2b. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in 'old-growth' forest #2, Lancaster, ON, (OG-2) ...... 60 Appendix 2.2e. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in 'old-growth' forest #3, Morrisburg, ON, (OG-3) ...... 64 Appendix 2.2d. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #1, Lancaster, ON, (MF-l) ...... 68 Appendix 2.2e. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #2, Bainsville, ON, (MF-2) ...... 72 Appendix 2.2f. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #3, Bouck's Hill/Williamsburg, ON, (MF-3) ...... 76 Appendix 3.1a. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #1, Cornwall, ON, (OG-l) ...... 111 Appendix 3.1b. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in 'old-growth' forest #2, Lancaster, ON, (00-2) ...... 113 Appendix 3.1e. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in 'old-growth' forest #3, Morrisburg (Upper Canada), ON, (OG-3) ...... 115

VIl Appendix 3.1d. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #1, Lancaster, ON, (MF-1) ...... 117 Appendix 3.Ie. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #2, Bainsville, ON, (MF-2) ...... 119 Appendix 3.H. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #3, Bouck's Hill (near Williamsburg), ON, (MF-3) ...... 121

V1l1 ACKNOWLEDGEMENTS 1 would like to thank my supervisors, Robert S. Anderson (Canadian Museum of Nature) and Terry A. Wheeler (Lyman Entomological Museum, McGill University), for their support and direction during the long arduous task of completing my Master's. In addition to editing manuscripts and answering pesky little questions, Bob provided financial support and Terry provided office and lab space throughout the study. 1 would also like to thank Christopher M. Buddle (Mc Gill University), a member of my advisory committee who provided valuable advice on project design and data analysis. 1 greatly appreciate the help provided by several taxonomists: Stewart B. Peck (Carleton University) identified and verified many leiodid specimens, Andy Cline (Louisiana State Arthropod Museum) identified and verified nitidulids, and Scott Redhead (National Mycological Herbarium, Agriculture and Agri-Food Canada), not only identified fungi for me, but accompanied me into the mosquito-ridden field. AIso, several specialists (Floyd Shockley, Andy Cline, and Paul Skelley) were kind enough to answer mye-mails and provide information on their respective taxa. While searching for and gaining permission to access my study sites, a number of people provided me with information and took me on site tours: Jim Hendry and Harry Hutchinson (Resource Stewardship Council of Stormont, Dundas & Glengarry), Martin Streit and Peter Wensink (Domtar Inc.), and Barry Hughes and Gerben Schaillee (St. Lawrence Parks Commission). 1 also thank the landowners who granted me access to their forests.

Financial support was provided through NSERC Di~covery grants to R.S. Anderson and T.A. Wheeler and a Nature Discovery Fund grant to R.M. Zeran. A big thank you to Larry Shaver for helping me ob tain materials for, construct, install and dismantle all of the flight-intercept traps. Thank you also to Norma Zeran and Bert Zeran for occasional forays into the (still mosquito-infested) forest. Finally, 1 would like to thank aU ofmy colleagues at the Lyman Entomological Museum (Stephanie Boucher, Scott Brooks, Eleanor Fast, Marjolaine Giroux, Jade Savage, Duncan Se1by and Terry Wheeler) for helping make the time pass, especiallyon Thursdays. A special thank you to Marjolaine for providing a French translation ofthe abstract and to Duncan for driving me places whenever 1 asked him to.

IX PREFACE This thesis is composed of four chapters, two of which are original manuscripts which will be submitted for publication in refereed journals.

Chapter 1 This chapter is a general introduction and literature review.

Chapter 2 This chapter is a manuscript in preparation for submission to Biodiversity and Conservation. Zeran R.M, Anderson R.S, and Wheeler T.A. Biodiversity offungivorous beetles (Coleoptera) in managed and 'old-growth' hemlock-hardwood forests in southeastern Ontario.

Chapter 3 This chapter is a manuscript in preparation for submission to The Canadian Entomologist. Zeran R.M., Anderson R.S, and Wheeler T.A. Sap beetles (Coleoptera: Nitidulidae) in managed and 'old-growth' forests in southeastern Ontario, Canada.

Chapter4 This chapter is a general conclusion.

x CONTRIBUTION OF AUTHORS R. Zeran designed the research for both manuscripts and also carried out field sampling, specimen preparation and identification, data analyses, and writing. R.S. Anderson supervised the research, provided financial support for the duration of the study and edited the manuscripts. T .A. Wheeler supervised the thesis research, provided laboratory facilities and materials and edited the manuscripts.

Xl ABSTRACT The species richness, abundance and composition of saproxylic Coleoptera was compared between 'old-growth' and mature-managed hemlock-hardwood forests in southeastem Ontario, Canada. Beetles were sampled weekly from 29 April until 3 October 2003 using large-area flight-intercept traps (FITs) and trunk-window traps (TTs). Analyses were conducted using the Fisher's ex and Simpson's diversity indices, rarefaction, indicator species analysis and c1uster analysis. A total of Il,888 fungivorous Coleoptera was collected from Il families and 73 species (exc1uding Nitidulidae). Nitidulidae were analysed separately with traps yielding 2,129 sap beetles comprising 30 species. The species richness and abundance of fungivorous Coleoptera did not differ significantly between the two forest types. Conversely, the species abundance of nitidulid beetles was higher in managed forests and the species richness higher in 'old growth' forests. Several species were strongly associated with either managed or 'old growth' forest types. Certain species such as Anisotoma inops (Leiodidae) and Glischrochilus sanguinolentus (Nitidulidae) were much more frequently caught in TTs than in FITs.

XIl RÉSUMÉ La richesse en espèces, l'abondance ainsi que la composition en coléoptères saproxyliques entre une « vieille» forêt et une forêt mature aménagée de pruche et de feuillus du sud-est de l'Ontario (Canada), ont été comparés. Les coléoptères ont été récoltés chaque semaine, du 29 avril au 3 octobre 2003, à l'aide d'un piège d'interception à grande surface et d'un piège-tronc à fenêtre. Des analyses, incluant les indices de diversité de Simpson et ex de Fisher, une courbe de raréfaction, la valeur indicatrice des espèces ainsi qu'une analyse de groupement, ont été effectuées. Au total, Il 888 coléoptères fongivores, appartenant à Il familles et 73 espèces (excluant les Nitidulidae), ont été capturés. Les analyses concernant les Nititulides ont été effectuées séparément puisque leur capture s'élevait à 2 129 spécimens appartenant à 30 espèces. Aucune différence significative de la richesse en espèces et de l'abondance des coléoptères fongivores n'a été notée entre les deux types de forêt. Inversement, l'abondance des espèces de Nitidulides fut supérieure dans forêt mature aménagée alors que la richesse en espèces fut plus grande dans la« vieille» forêt. Plusieurs espèces étaient fortement associées soit à la forêt mature aménagée, soit à la « vieille» forêt. Certaines espèces telles Anisotoma inops (Leiodidae) et Glischrochilus sanguinolentus (Nitidulidae) furent capturés plus fréquemment dans le piège-tronc à fenêtre que dans le piège d'interception à grande surface.

Xlll CHAPTER 1 - Introduction and Literature Review

"There is something in the ponderous stillness ofthese forests - something in their wild, torn, mossy darkness, their utter solitude and mournful silence which impresses the traveUer with a new aspect each time he sees them .. .ln Upper Canada the endless hills of pine give way at last, or at most stand thinly intermingled with gigantic beeches, taU hemlocks and ash, with maples, birch and wild sycamore, the underwood ofthese great leafy hills. Mile after mile, and hour after hour ofsuch a route was passed - a dark black solitude, with here and there a vista opening up, showing the massive trunks, grey as cathedral ruins, which bore the rich canopy ofleaves aloft." An early settler to eastem Ontario, cited in Keddy (1994)

General Introduction Biodiversity refers to the variety of ecosystems and habitats, the number and variety of species within them, and the genetic variability within each species' population (Caldecott et al. 1996). Since its use in the title of a 1988 book edited by E.O. Wilson, the term 'biodiversity' has been used with increasing frequency and has been attributed an array of different meanings (Haila and Kouki 1994). In ecology, biodiversity (or biological diversity) consists oftwo main components: the variety (or richness) ofspecies and the relative abundance of species (Magurran 2004). Over the past decade, interest in the study ofbiodiversity has increased, especially in ecosystems such as forests. By far, the most abundant multicellular organisms in forests are arthropods. Arthropods currently constitute approximately 64% of global biodiversity (Finnamore 1996), and estimates of species still to be discovered raise this figure significantly. Insects can be useful bioindicators because large sample sizes can be obtained with ease and minimal bias, they have relative1y short generation times, and they are frequently sensitive to local habitat changes (McGeogh 1998). Coleoptera, in particular, are useful in biodiversity research and environmental assessment for a number of reasons: their taxonomy and distributions are relatively well known in Canada; they are taxonomically and trophically diverse; theyare easy to preserve and prepare for subsequent identification; and many previous studies have focused on Coleoptera and the new data can be compared with that from existing studies (Hammond 1997). Coleoptera

1 are aiso one of the most diverse groups offorest insects and account for approximately 20-40% of total arthropod diversity (Carlton and Robison 1998, Grove 2002). And, of particular relevance herein, they represent a large portion of the groups found in dead wood. Ecological monitoring and other comparative studies have traditionally been carried out using megafauna and megaflora, large 'visible' organisms such as vascular plants and vertebrates. However, the less visible micro-flora and -fauna (such as arthropods and fungi) make up 95% of total diversity and drive most ecosystem processes (Finnamore 1996). Recognizing the importance ofthese organisms, an increasing amount of work has recently been carried out to monitor and evaluate the status of the microfauna/flora of ecosystems such as forests. Such studies have become increasingly important as we realize the effects ofhuman activity on various habitats and environments. However, the majority ofthis work has taken place in Fennoscandia and, for most other areas in the world, such information is sti11lacking. This situation must be rectified if we are to fully understand, manage and protect areas under development and/or subject to human-induced stress.

Eastern Ontario Forests The Great Lakes-St. Lawrence forest region (Rowe 1972) is a transitional zone between the boreal and deciduous forests and constitutes approximately 22% of Ontario's forest coyer. Between the mid-1700s and early 1900s an influx of European settlers resulted in the destruction of a large portion of Ontario' s temperate mixed-wood old growth forest (Keddy 1994, Anonymous 1999). Forest coyer in eastem Ontario is currently estimated at about 38 percent ofwhat it was historically (Anonymous 1997). Over 95% of the habitat in this region has been lost to development; remaining habitat mostly consists of wetlands and abandoned farmlands undergoing reforestation. The area's forests are highly fragmented with little connectivity between patches and little interior habitat in remaining stands. The region is heavily populated and natural areas have been greatly modified by agriculture and urbanization, making conservation of the area's remaining old growth forests aIl the more important.

2 The pre-settlement forests of the Great Lakes-St.Lawrence Forest Region (GLSL) were dominated by sugar maple (Acer saccharum Marsh.), beech (Fagus grandifolia Ehrh.), and hemlock (Tsuga Canadensis (L.) Carr), with lesser numbers of white pine (Pin us strobes L.), elm (Ulmus Americana L.), and basswood (Tilia Americana L.) (Keddy 1994, Anonymous 1997, 1999). Hemlock and beech are much less noticeable in our forests today. Hemlock was decimated by clearance for agriculture, by logging (railway ties), and by potash and tanbark production (Suffling et al. 2003). Similarly, beech, considered less valuable, was cleared from productive soils and shipped as potash (Suffling et al. 2003). Both hemlock and beech have failed to re-establish over large areas, attributed at times to large deer populations and beech bark disease (Suffling et al. 2003). This is unfortunate since, in forest landscapes, hemlock can reach ages greater than any other tree species in the GLSL; and beech, when dead or moribund, provides habitat for a remarkably diverse array of species.

Old-growth forests It is difficult to develop a precise definition for old-growth because old-growth characteristics will vary from one forest type to another; differences in soils, climate, physical geology, and disturbance regimes, make an all-encompassing definition of old growth virtually impossible. Generally, scientists, environmentalists and foresters recognize old-growth as being a forest that is relatively old (where a significant proportion oftrees have exceeded their average life-span) and relatively undisturbed (i.e., free of severe human disturbances such as logging) (Stabb 1996). Old-growth forests are also evaluated based on the number and quality of 'old-growth characteristics' that they display. These characteristics (White 1990, McCarthy 1995, Anonymous 1996, 1999, Stabb 1996, Uhlig et al. 2001) are discussed below. a. Dominant trees are large and old. This is perhaps the easiest way in which the casual observer can recognize old-growth. The minimum age of an old-growth forest is usually one-half the average life-span of the dominant trees in the forest. Thus, the boreal forest of Canada usually reaches old-growth age at just over 90-100 years, the Great Lakes-St. Lawrence forest at about 150-200 years, and the western coastal rainforests of British Columbia at 200-300 years. It is important to realize that the age of

3 the tree does not always translate into a large diameter for the tree; tree growth depends greatly on various aspects of site quality such as soil profile, microc1imate and surrounding trees. b. All-aged or uneven-aged stand structure. The forest is characterized by trees having a range of ages and diameters. c. Little or no sign of human disturbance. This is potentially one of the most controversial characteristics of old-growth. In areas ofhigh human settlement and development, there is little to no forest remaining that has never been influenced by humans in sorne way. Sorne argue that any forest disturbed by humans at sorne time is not old-growth, while others recognize old-growth as forests which have not been severely disturbed for a long time. It is simpler to divide the terminology: a pristine or virgin forest is one which has never been influenced by humans; while an old-growth forest is one which has developed naturally for a long time (80-100 years) without ever experiencing a major human-caused disturbance (such as c1ear-cut logging or large-scale harvest). d. Multi-Iayered canopy structure. This inc1udes: super-canopy trees such as taU white pines that poke up through the canopy and provide nesting, resting and refuge areas for wildlife; canopy trees that form a continuous ceiling and shade layers below; understory trees (small trees growing beneath the canopy); and shrubs and saplings. e. Coarse woody debris. In recent years a vast amount of literature has been published that describes the importance of coarse woody debris to the forest ecosystem. This characteristic will be discussed in a following section. f. Snags and cavity trees. Snags are standing dead trees that provide habitat for many forest wildlife species and are an integral part of the forest ecosystem. One ofthe most characteristic aspects of snags is the presence of numerous cavities (holes excavated by animaIs for feeding, nesting, resting, or escape). Cavities may also be present in living trees. g. Natural tree-fall gaps. Old-growth forests typically display many small open areas where the forest canopy is disrupted, these are termed 'gaps'. Usually the result of high winds, these gaps are created when one to several trees are blown down, opening an

4 area of the forest to increased sunlight penetration, and allowing regeneration and succession to occur. h. Pit and mound topography. Pits are formed when a tree is uprooted by wind and falls. Its root mat and associated soil are thus ripped from the forest floor, creating a pit-like depression. The mound forms as the root mat decays. Pits and mounds are important in forest nutrient cycling and understory diversity and are generally absent from or greatly reduced in human-disturbed forests. Forests which have been c1eared for agriculture or harvested repeatedly will not usually contain a sculpted pit and mound topography. Old-growth forests provide essential habitat for numerous plant and wildlife species. Birds, such as the Barred Owl (Strix varia) and Pileated Woodpecker (Dryocopus pileatus), use natural cavities and snags for resting and feeding. Many other birds require the habitat provided by mature forest (e.g., Red-shouldered Hawk, Buteo lineatus; Scarlet Tanager, Piranga olivacea; Wood Thrush, Hylocïchla mustelina; Brown Creeper, Certhia americana). Bats, such as the silver-haired bat, Lasionycteris noctivagans, roost in snags. Old-growth forest is also important habitat for redback voles (Clethrionomys gapperi), martens (Martes americana), red-backed salamanders (Plethodon cinereus), spotted salamanders (Ambystoma maculatum), and ring-necked snakes (Diadophis punctatus)(Anonymous 1999, Bellhouse and Naylor 1996). Over the past few decades a large volume of research has been conducted on the importance of old-growth forests and old-growth forest characteristics to invertebrates (Siitonen and Martikainen 1994, Kaila et al. 1997, Nilsson and Baranowski 1997, Martikainen et al. 2000). The relationship between old growth forest and invertebrates will be discussed in more detail in subsequent sections.

Downed woody debris Downed woody debris (DWD) can be divided into two categories: fine woody debris (FWD) and coarse woody debris (CWD). FWD is usually considered to be dead and downed branches, twigs and small tree or shrub bol es that are under 7.5 cm in diameter (Bellhouse and Naylor 1996). However, the limits ofFWD vary from project to project and from organization to organization and wood up to > 15 cm diameter is

5 sometimes considered FWD. Coarse woody debris inc1udes downed and dead tree and shrub boles, large limbs and other large woody pieces, standing dead trees leaning more than 45° degrees from vertical, and sometimes even cut stumps. As with FWD, the lower diameter limit of CWD varies with the study, but it is fairly safe to c1assify any dead downed wood over 10 cm in diameter as CWD. In addition to size, one of the most frequently measured characteristics ofDWD is the stage of decay. Decay stage classification varies from one region to another (sorne areas only inc1ude three stages of decay, while others have as many as seven stages). The most common and widely accepted classification system (at least in the temperate forests ofeastern Ontario) is based on five decay stages (e.g., Bellhouse and Naylor 1996). Decay class is based on the presence or absence oftwigs and bark, the texture of the bark, the shape ofthe log, the colour of the wood, and the amount of the log that is touching the ground (Bellhouse and Naylor 1996). The rate of decay ofDWD depends on a variety of factors: (1) when and how the wood fell (windfall, senescence, snags, cutting); (2) the size of the wood (large diameter logs decay more slowly); (3) the species oftree (coniferous trees generally decay more slowly than deciduous trees); and (4) c1imate and microclimate at the site (e.g., temperature and rainfall) (Bellhouse and Naylor 1996). The rate at which a log decays is important because its decay stage will determine how and what animaIs, plants, and fungi use the wood. Downed woody debris is important to the forest ecosystem; it influences nutrient cyc1ing, water retenti on, soil formation and erosion (Harmon et al. 1986, Bellhouse and Naylor 1996). It also provides a seedbed for new tree growth and provides critical habitat for a variety ofwildlife and arthropod species (Harmon et al. 1986, Bellhouse and Naylor 1996). DWD is also important in streams as fish habitat (Harmon et al. 1986). In Ontario, 89 vertebrate species use DWD as coyer (restinglroosting, escape, travel), for display sites, for feeding (seeking, caching, butchering), for hibernation or as sites for reproduction (nesting) (Bellhouse and Naylor 1996). The number of invertebrates who depend on the presence of dead wood for their survival is vastly greater. The functional importance ofDWD to invertebrates depends on the amount of dead wood, its size distribution, its spatial arrangement at the site, its decay stage, and its

6 position (i.e., raised versus ground contact) (Hannon et al. 1986). For example, wood boring beetles (e.g., Scolytidae) are frequently found in newly dead wood (and less often in significantly decayed wood) and may 'precondition' the wood for a number of succeeding species (Graham 1925, Harmon et al. 1986, Speight 1989, Hammond et al. 2001). Invertebrates may use DWD for a variety ofreasons: as protection from the environment (e.g., shelter for overwintering insects), as a food source (both direct and indirect), as a hibernation site, and as a nesting and/or breeding site (Harmon et al. 1986; Hammond 1997). The dependence of invertebrates on DWD varies from one species to another; sorne use the wood only in passing, while others could not survive without a continuous supply ofDWD. Many studies have shown that the presence of dead wood in large quantities is extremely important to the survival ofsaproxylic beetles (e.g., Siitonen 1994, 0kland et al. 1996, NiemeHi 1997, Scheigg 2000, Martikainen et al. 2000). This point will be discussed further in the following section. Dead wood is also very important for the species diversity ofwood-inhabiting fungi (Lindblad 1998, Grove 2002). Logs in later stages of decomposition increase the diversity ofwood-inhabiting fungi, and may act as sources for fungal species colonizing freshly fallen dead wood (Lindblad 1998). Bader et al. (1995) suggested that three main factors influence the species composition ofwood-inhabiting fungi in forest stands: (1) the abundance oflogs - more CWD provides more substrate for fungi to colonize; (2) log size can influence the total number of species per log and the species composition - larger logs have more surface area and can support more fungal mycelia and a greater diversity of fungal species; (3) decay stages and rate of decay oflogs (influenced by moi sture, temperature and light conditions of site) - a more diverse log decay range will support a more diverse array of fungal species. FWD is also an important substratum influencing the diversity ofwood-inhabiting fungi, especially ascomycetes. In Sweden, Norden et al. (2004) found that ascomycetes were found predominately on FWD (75%) and basidiomycetes were found mainly on CWD (although 30% ofbasidiomycetes were also found on FWD). Snags are sometimes considered as DWD in inventories. Snags provide much of the same function in the forest ecosystem as do downed logs and are the primary source of nesting cavities for birds and wildlife. However, snags can be quite different from

7 downed wood in terms of abiotic factors such as moisture content and light exposure, and these differences can greatly influence the insect fauna using snags. In Norway, Sverdrup-Thygeson and Ims (2002) found large differences in the beetle composition between snags and downed logs. Snags had higher species richness and were more likely to harbour red-listed (threatened) species. They hypothesized that this may occur because sun exposure might have a stronger effect on the beetle species composition in snags than in logs. A natural forest ecosystem will generally have a continuous input ofDWD from mortalityagents such as wind, tire, insect damage, disease, and natural suppression and competition oftrees (Harmon et al. 1986). Forest management can alter these processes, changing the dynamics of dead wood and ofthe fungi, insects, and vertebrate communities that depend upon this wood.

Saproxylic beetles and forest management Saproxylic beetles are beetles that depend, during sorne part of their life-cycle, on the wood ofmoribund or dead trees, on wood-inhabiting fungi, or upon the presence of other saproxylic species (Speight 1989). As noted above, dead wood is an extremely important resource to saproxylic beetles (Siitonen 1994, 0kland et al. 1996, Niemela 1997, Jonsell et al. 1998, Scheigg 2000, Martikainen et al. 2000). Thus, removing or reducing the volume ofDWD from forests (i.e. during harvesting operations) can seriously affect saproxylic insect fauna. In natural unmanaged forests, trees dying from natural disturbances (e.g., wind, tire) are the primary source ofCWD. In older natural forests mortality caused by tree senescence and small-scale gap disturbances are important and increase with increasing stand age. Thinning and cutting a forest can truncate these processes by removing large trees before they die (Hansen et al. 1991, Siitonen et al. 2000). There are many types of forest management practices that have been studied in relation to saproxylic beetles, but the two most common practices are selective cutting and clear-cutting. In general, forest management can affect saproxylic beetles in several ways: (1) by changing the tree-species composition and stand structure and thus, changing the composition of dead wood (and fungus) available for breeding; (2) by

8 affecting the mortality rate of live trees and thus the dynamics of dead wood; (3) by changing the within-stand microclimate; and (4) by affecting the interactions among species (Martikainen et al. 1999). Much research has been devoted to understanding the connection between volume of dead wood and forest condition. In most studies, dead wood volumes were highest in old-growth stands and lower in managed stands (Goodburn and Lorimer 1998, Siitonen et al. 2000, Martikainen et al. 2000). Sometimes the difference in volume is drastic: for example, Siitonen et al. (2000) found that the average volume of coarse woody debris was 14 m3/ha in mature-managed forests, 22 m3/ha in overmature-managed forests and 111 m3/ha in old-growth stands. Volume is not the only characteristic of dead wood affected by forest management. In Finland, Simila et al. (2003) found that, due to intensive forest management, the volume of dead wood had decreased and its quality (e.g., diameter and decay stage) was less diverse in managed forests versus older undisturbed forests. Similarly, Siitonen et al. (2000) found that most ofthe volume of CWD in old-growth forests belonged to logs in large diameter classes (20cm to +40cm), while in managed forests, most ofthe CWD volume belonged to small diameter classes (5-l9cm). Gore and Patterson (1986) found similar results in New Hampshire. In Sweden, Bader et al. (1995) noted that forest cutting can result in "interruptions in the continuity of decaying logs", causing fewer logs in higher decay stages to be present in intensely-managed forests. The range of decay stages of dead wood can also have a significant influence on the saproxylic insect fauna (Grove 2002); while sorne species colonize freshly dead wood, others utilize only CWD in mid and/or late stages of decay. Over the past few decades, many studies have attempted to better understand the dynamics between dead wood and saproxylic arthropods in forest ecosystems in order to better conserve total forest biodiversity. Martikainen et al. (1999,2000) found that the species richness of saproxylic beetles was significantly higher in old-growth than in managed forests due mainly to the greater amount and diversity of decaying wood in old growth forests. 0kland et al. (1996) found that past extensive forest cutting was negatively correlated with the species richness of saproxylic beetles and suggested that this occurred because of decreasing recolonization of the beetles into the eut forest due to

9 increasing distance from a source habitat. Nilsson and Baranowski (1997) found that saproxylie beetles living in hollow beeeh trees had lower species richness in managed stands than in near-primeval stands. Siitonen and Martikainen (1994), in a study ofthe beetles in Russian and Finnish Karelia, found that rare saproxylic beetle species were more common in forests with less management and a higher volume of dead wood. Hammond et al. (2001) found that species diversity of saproxylie beetles was higher in old rather than in mature forest stands. Although saproxylic beetle species inhabit all kinds of forests, the faunal composition sometimes differs drastically in managed forests from that of old-growth forests. Siitonen (1994) found that specialist beetle species were most abundant in forest with a large supply of decaying wood, while generalist species did not prefer one site over the other. Kaila et al. (1997) studied the beetle fauna inhabiting dead standing birch trees in clear-cuts and mature forest. They found that distinct beetle assemblages were associated with each different habitat (e.g., species associated with open areas were found predominantly in clear cut areas, while species assoeiated with closed forest occurred mainly in the mature forest areas). Vaisanen et al. (1993) studied the sub-eortical beetle fauna in Finland and found that species richness and abundance tended to be higher in managed forests than in primeval forest; however, several species were only found in one or the other forest type, with rare species being more common in primeval stands. Berg et al. (1994) noted that threatened species in Sweden are more commonly found in old forests than in managed forests. Decaying wood in the form of stumps and logging slash are eommon in managed forests and provide habitat for certain saproxylic beetles. However, the laek of abundant deeaying snags and logs in managed forest can be detrimental to other species that specialize solely on such wood. Perhaps the most studied group ofbeetles in forest management studies is the Carabidae (ground beetles). Werner and Raffa (2000) found that several species of carabid beetles in eastern hemlock forests showed specifie habitat associations, e.g., sorne preferred old-growth sites over managed and vice versa. Several other studies have reached similar conclusions (e.g., Niemela et al. 1988, Niemela et al. 1992, Spence et al. 1996, Heliola et al. 2001, Koivula et al. 2002). For earabid beetles, the most common trends following forest cutting have been (1) the species common to open habitats

10 increase, (2) there is an initial decline of forest generalists, and (3) forest specialists disappear (Niemela et al. 1993, Werner and Raffa 2000, Heliola et al. 2001, Koivula et al. 2002). However, carabid beetles may not be the best beetles to demonstrate the effect of forest management practices and, although they may be useful bioindicators, it is still unclear how weIl they represent the responses of other species (Raino and Niemela 2003). The above-mentioned studies deal only with beetles; however, work on other groups of arthropods has been conducted with much the same result. Coyle (1981) found that clearcutting reduced the diversity of web-building spiders, but increased or had no effect on the numbers of hunting spiders. He postulated that removal of the forest canopy results in microclimate changes that are not preferred by web-building spiders. 0kland (1994) studied Mycetophilidae (Diptera) in Norway and found that semi-natural forests were more species-rich and contained more rare species than clear cuts and managed forests. Other examples include work by Punttila et al. (1994) on Finnish ants (Hymenoptera: Formicidae), by Huhta (1976) on soil invertebrates in Finland and by Buddle et al. (2000) on spiders. Much of the work carried out on saproxylic fauna has provided suggestions on the best management practices for forests today. Without a doubt, undisturbed old-growth forest must be set aside to sustain specialist species and to serve as sources of recolonization for managed forests (Niemela 1997). But what of managed forests? Several studies have suggested that leaving dead and dying wood behind during forest thinning and clear-cutting could greatly aid in preserving diversity in managed forests (Kaila et al. 1994, Kaila et al. 1997, Schiegg, 2000, Jonsell and Weslien 2003, Sverdrup Thygeson and Ims 2002). Applying harvesting methods to retain old-growth characteristics (e.g., leaving large living trees, snags and logs behind after logging) will increase the structural diversity and old-growth attributes in managed forests (Siitonen et al. 2000). Increasing the total volume of dead wood in managed forests is important; however, increasing the diversity and continuity (e.g., a variety of size and decay classes) ofthat dead wood is just as important (Bader et al. 1995; Simila et al. 2003). Most studies agree that in order to maintain a diversity of saproxylic invertebrate species we

11 need to ensure that a variety of substrate types and management methods are preserved (Jonsell et al. 1998; Werner and Raffa 2000).

Fungus-insect relationships Wood-decaying fungi are an exceptionally variable microhabitat and resource of hundreds of species of insects. The majority of fungivorous beetles inhabit a few closely related fungal species (Komonen 2001,2003). Several beetle species, however, can be associated with only one fungal hosto Kaila et al. (1994) found that distinct beetle assemblages were associated with different polypore species. Even taxonomically and ecologically related fungi can host completely different insect fauna and this can make the insect species more vulnerable to forest disturbance (Komonen 2001). Fungus-inhabiting beetles differ in their fungal substrate preferences; factors that affect these choices include successional stage, degree of sun exposure, height above ground, and fruiting body size (Jonsell and Nordlander 2002). Hagvar (1999) suggested that saproxylic beetles seek out living sporocarps either for feedinglbreeding, or by following the odour of living sporocarps in order to find dead wood in which to breed. Beetles may be beneficial to fungus ifthey remove mites and larvae that block fungal pores and ifthey act as vectors, spreading spores to other logs and bark (Hagvar 1999). Most fungal species are very cryptic, existing merely as mycelia in wood or may be seen only in small, short-lived fruiting bodies (Jonsell and Weslien 2003). There is sorne indication that there may be close relationships between saproxylic beetles living inside wood and the fungus present in wood only as these inconspicuous mycelia (Jonsell and Weslien 2003). Thus, the number ofinvertebrates dependent on fungi is probably underestimated because species living on mycelia in the wood are difficult to distinguish from those living on the wood alone (Jonsell et al. 1998, Komonen 2003). Many studies have found that the diversity ofwood-inhabiting fungi may be a good indicator ofspecies richness ofmany saproxylic beetle species (e.g., Kaila et al. 1994, 0kland et al. 1996, Komonen et al. 2000, Komonen 2001). Not all studies have found, however, that wood-inhabiting fungus is directly related to the species richness of certain saproxylic beetles. For example, Sverdrup-Thygeson (2001) found that there was

12 no significant correlation between wood-rotting fungus and the species richness ofred listed saproxylic beetles in Norway. The diversity ofwood-inhabiting fungi can also be negatively affected by forest management (e.g., Penttila et al. 2004) and, this may reduce the abundance of certain invertebrates that depend on the fungi (Bader et al. 1995, Komonen et al. 2000). Sorne fungus-inhabiting insects that are monophagous on certain species of fungus (microhabitat), also have specific macrohabitat requirements (e.g., they are restricted to old-growth forests even though their fungal host may also occur in managed forests) (Komonen et al. 2000, Jonsell and Nordlander 2002). Many variables associated with saproxylic beetles are interconnected, e.g., volume of dead wood and number oflogs in large diameter classes (0kland et al. 1996) and the abundance ofwood-inhabiting fungi. Thunes et al. (2000), in a study ofbeetles inhabiting bracket fungus in Norway, found that the main factor influencing beetle diversity was the volume of dead wood in the vicinity of the sampling site due to the higher number of species per unit volume of sporocarps in areas with high levels of dead wood. Such areas also hosted more red-listed beetle species. As Hammond et al. (2001) noted, changes in the beetle fauna inhabiting CWD could be tightly linked to changes in the fungal community structure over time.

13 OBJECTIVES The first objective ofthis study was to conduct a faunal inventory and to document the diversity and distribution patterns of saproxylic beetles in the remnant forests ofsoutheastern Ontario. It will provide updated knowledge of the current distributions of many species as weIl as provide baseline information for future monitoring, research or educational work in the area. There were two components to this project: the first was to inventory fungivorous Coleoptera from the Leiodidae (Leiodinae), Erotylidae, Endomychidae, Mycetophagidae and other less frequently encountered families (chapter 2). The second component was to conduct an inventory of the Nitidulidae (chapter 3). Nitidulids have a wide range offeeding strategies, are not considered strict fungivores and, apart from the species that are considered minor agricultural pests, little is known about the ecology and distribution of the majority of sap beetles in eastern Ontario. Phenological patterns of selected fungivorous and nitidulid species were also examined and, where possible, compared to published literature. The second objective was to compare the species richness, species composition and relative abundance offungus-beetle (chapter 2) and nitidulid beetle (chapter 3) assemblages in old-growth hemlock-hardwood forests to the fauna from managed forests of similar age and tree species composition. While a large quantity of such research has recently been completed in Fennoscandia and in western Canada, very little work has occurred in the Great Lakes-St. Lawrence Forest region. It is important to examine the effects that forest management may have on smaller organisms like insects, since they make up the majority of species using the forest and are responsible for a variety of essential ecosystem functions (e.g., decomposition and pollination).

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18 0kland B, Bakke A, Hagvar Sand Kvamme T. 1996. What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southern Norway. Biodiversity and Conservation 5: 75-100. Penttila R, Siitonen J and Kuusinen M. 2004. Polypore diversity in managed and old growth boreal Picea abies forests in southern Finland. Biological Conservation 117: 271-283. Punttila P, Haila Y, Niemela J and Pajunen T. 1994. Ant communities in fragments of old-growth taiga and managed surroundings. Annales Zoologici Fennici 31: 131- 144. Raino J and Niemela J. 2003. Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation 12: 487-506. Rowe JS. 1972. Forest Regions of Canada. Department ofFisheries and the Environment. Canadian Forestry Service, Publication No. 1300. 172pp. Schiegg K. 2000. Are there saproxylic beetle species characteristic ofhigh dead wood connectivity? Ecography 23: 579-587. Siitonen J. 1994. Decaying wood and saproxylic Coleoptera in two old spruce forests: a comparison based on two sampling methods. Annales Zoologici Fennici 31: 89- 95. Siitonen J and Martikainen P. 1994. Occurrence ofrare and threatened insects living on decaying Populus tremula: A comparison between Finnish and Russian Karelia. Scandinavian Journal of Forest Research 9: 185-191. Siitonen J, Martikainen P, Puntilla P and Rauh J. 2000. Coarse woody debris and stand characteristics in mature managed and old-growth boreal mesic forests in southern Finland. Forest Ecology and Management 128: 211-225. Simila M, Kouki J and Martikainen P. 2003. Saproxylic beetles in managed and seminatural Scots pine forests: quality of dead wood matters. Forest Ecology and Management 174: 365-381. Speight MCD. 1989. Saproxylic invertebrates and their conservation. Councilof Europe, Strasbourg.

19 Spence JR, Langor DW, Niemelâ J, Carcamo H and Cameron R. 1996. Northern forestry and carabids: the case for concern about old-growth species. Annales Zoologici Fennici 33: 173-184. Stabb M. 1996. Ontario's old growth: a leamer's handbook. Canadian Nature Federation, Ottawa, ON, 47 p. Suffling R, Evans M and Perara A. 2003. Presettlement forest in southern Ontario: ecosystems measured through a cultural prism. The Forestry Chronic1e 79: 485- 501. Sverdrup-Thygeson A. 2001. Can 'continuity indicator species' predict species richness or red-listed species of saproxylic beetles? Biodiversity and Conservation 10: 815-832. Sverdrup-Thygeson A and Ims R.A. 2002. The effect of forest c1earcutting in Norway on the community ofsaproxylic beetles on aspen. Biological Conservation 106: 347- 357. Thunes KR, Midtgaard F and Gjerde 1. 2000. Diversity ofColeoptera of the bracket fungus Fomitopsis pinicola in a Norwegian spruce forest. Biodiversityand Conservation 9: 833-852. Uhlig P, Harris A, Craig G, Bowling C, Chambers B, Naylor B, and Beemer G. 2001. Old growth forest definitions for Ontario. Ontario Ministry ofNatural Resources, Queen's Printer for Ontario, Toronto, ON. 53pp. Vâisanen R, Bistrom 0, and Heliovaara K. 1993. Sub-cortical Coleoptera in dead pines and spruces: is primeval species composition maintained in managed forests? Biodiversity and Conservation 2: 95-113. Werner SM and Raffa KF. 2000. Effects offorest management practices on the diversity of ground-occurring beetles in mixed northern hardwood forests of the Great Lakes Region. Forest Ecology and Management 139: 135-155. White Dl 1990. Preliminary Definitions and Evaluation of Old-Growth Forest in Eastern Ontario. Ontario Ministry ofNatural Resources, Kemptville, Ontario.

20 CONNECTING STATEMENT While many comparative biodiversity studies on Coleoptera have occurred in northem Europe, very few have taken place in North America, especially in the Great Lakes-St. Lawrence Region of Canada. This was the first season-Iong biodiversity study in eastem Ontario focusing specifically on fungivorous Coleoptera. Data from this study will provide valuable information for future monitoring or inventory work in the area and will provide updated distribution data for many Coleopteran species. Over the past few decades, comparative arthropod studies investigating the impacts of forest management have occurred across the globe. As Chapter 1 showed, old-growth forests and old-growth forest characteristics can be extremely important to the survival ofmany species. Studies such as this will provide information to forest managers and conservationists on the impact of forest harvest operations and can help to ensure that aIl components of the forest ecosystem remain intact for future generations.

21 CHAPTER 2 - Biodiversity of fungivorous beetles (Coleoptera) in managed and 'old-growth' hemlock-hardwood forests in southeastern Ontario

ABSTRACT Fungivorous Coleoptera were sampled from 'old-growth' and mature-managed hemlock-hardwood forests in southeastem Ontario. Large-area flight-intercept traps (FITs) and trunk-window traps (TTs) were operated for 22 weeks in 2003, and yielded a total of Il,888 fungivorous beetles from Il families and comprising 73 species. The round fungus beetles (Leiodidae: Leiodinae) made up the bulk ofthis number with 10,386 individuals from 38 species. While 'old-growth' stands had higher volumes of coarse woody debris, species richness and abundance of fungivorous Coleoptera was similar between forest types, suggesting that past forest management did not have a significant effect on beetles in this study. Triplax macra LeConte was strongly associated with 'old-growth' forest, while Anisotoma blanchardi (Hom), Anogdus obsoletus (Melsheimer), Mycetina perpulchra (Newman) and a species of Agathidium showed significant associations with managed forests. Both Anisotoma inops WJ Brown and Triplax thoracica Say were collected far more frequently in TTs than in FIT s, illustrating the importance ofusing multiple trap types when sampling entire communities.

22 INTRODUCTION Anthropogenic disturbances such as logging, agriculture, fire suppression and urban development are responsible for the loss and fragmentation of forests worldwide. Most of Ontario's old-growth forests in the Great Lakes-St. Lawrence region (GLSL) were destroyed by logging, fires and settlement between the mid-1700s and the early 1900s (Suffling et al. 2003). A large portion ofCanada's economy still depends on timber harvest and agriculture (Anonymous 2001) making it important to manage existing forest stands for old-growth characteristics, not just for the conservation of large, visible species, but also for smaller organisms such as arthropods. In recent years, a large volume ofwork has been published on the diversity of lichens and bryophytes (e.g., Soderstrom 1988), fungi (e.g., Bader et al. 1995), and various arthropods, including spiders (e.g., Buddle et al. 2000), ants (e.g., Punttila et al. 1994), flies (e.g., 0kland 1994), beetles (e.g., Martikainen et al. 2000) and soil invertebrates (e.g., Huhta 1976) in managed and unmanaged forests. Much ofthis research has been carried out in Fennoscandia, with relatively few studies taking place in the GLSL forests of Canada and the United States (but see Chandler 1991; Chandler and Peck 1992; Werner and Raffa 2000). Saproxylic Coleoptera are one of the most diverse groups offorest insects (Speight 1989; Grove 2002). They depend, during sorne part of their life cycle, on the wood of moribund or dead trees, on wood-inhabiting fungi, or upon the presence of other saproxylic species (Speight 1989). Several factors have been shown to be important for saproxylic beetles, including the volume of dead wood (Martikainen et al. 1999; Schiegg 2000; Simila et al. 2003); the species, decay class and size class of dead wood (0kland et al. 1996; Jonsell et al. 1998; Simila et al. 2003); the abundance and diversity ofwood inhabiting fungi (0kland et al. 1996; Komonen 2003); the level of forest/site disturbance (Vliisiinen et al. 1993; Siitonen and Martikainen 1994); and landscape ecology and vegetational structure (0kland et al. 1996; Jokimaki et al. 1998). Numerous studies have shown that old-growth forests host several rare and/or specialist beetle species (Vliisiinen et al. 1993; Siitonen and Martikainen 1994; Berg et al. 1994). Old-growth forests have higher densities of snags and dead wood in larger diameter classes (Goodbum and Lorimer 1998; Siitonen et al. 2000; Simila et al. 2003)

23 and fungi are more diverse on coarse woody debris (CWD) with large diameters and in mid to late stages of decay (Bader et al. 1995). Because decaying wood and wood inhabiting fungi affect the diversity of saproxylic beetles (0kland et al. 1996; Thunes et al. 2000), one would expect that in old-growth forests, with large quantities of CWD that support large numbers ofwood-inhabiting fungi, fungivorous saproxylic beetles would be more diverse and abundant. Many studies of saproxylic Coleoptera have focused mainly on economically important bark-feeding or wood-boring species (such as members of the families Scolytidae, Cerambyeidae and Buprestidae), and few have dealt with speeies that are mainly fungivorous (Komonen 2003). Fungus-associated beetles dominate the forest landseape and, while not generally of large economic or social import, they are an important part of forest eeosystem proeesses, feeding on and dispersing fungi, which in turn are responsible for the majority ofwood deeomposition. The response of the Canadian saproxylie fauna to forest management praetices is not well known (Hammond 1997) and the few studies that have been published have foeused on the boreal forest (e.g., Spence et al. 1997; Hammond et al. 2001,2004). While the boreal forest is Canada's largest forest ecosystem, other areas, like the Great Lakes-St. Lawrence forest, are also important and have played a significant role in Canada's culture and history. Over the past decade there has been increased interest in forest inseet-fungus interactions; however, the majority ofthis work (as with work on saproxylic beetles in general) has been condueted in Fennoseandia (Komonen 2003). This study was earried out in the southem portion of eastem Ontario, Canada, an area dominated by agriculture and urbanization. The objectives ofthis study were to: (1) conduet a faunal inventory and document the diversity, distribution and phenology of fungivorous beetles in remnant forests of southeastem Ontario, and (2) compare the speeies richness, relative abundanee and speeies composition of selected fungus-beetle assemblages in 'old-growth' hemlock hardwood forests of southeastem Ontario to the fauna from managed forests of similar age and composition. 1 prediet that, beeause 'old-growth' forests have more mierohabitats with a greater variety of CWD and wood-inhabiting fungi, the diversity of fungivorous beetles is higher in 'old-growth' forests.

24 MATERIALS AND METHODS

Study Sites Six forest stands were selected in southeastem Ontario (Table 2.1, Fig. 2.1) in section L.2 of the Great Lakes-St. Lawrence Forest Region (Rowe 1972). Three forests were managed (MF-l, MF-2, MF-3) and three were considered 'old-growth' (OG-l, OG- 2,OG-3). For this study 'old-growth' forests were considered to be those which were of sufficient age (averaging over 120 years ofage), demonstrate many 'old-growth characteristics' (e.g., pit and mound topography, large trees, large volumes of coarse woody debris, multi-aged stand structure), and have remained mostly free ofhuman disturbance for at least several decades (e.g., Stabb 1996). Managed forests also demonstrated many 'old-growth' characteristics, only having slighter lower volumes of coarse woody debris and a less distinct pit and mound relief. AIl six forest stands were relatively smaIl (between 5.9 ha and 13.8 ha), fragmented remnant forests, with similar elevation, topography, soil, and c1imate characteristics. AIl sites were dominated by eastem hemlock (Tsuga canadensis (L.) Carr), with lesser quantities of sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifoUa Ehrh.), white pine (Pin us strobus L.) and a variety of other hardwoods. Managed forests ranged in age from 100- 150 years of age, while 'old-growth' forests were aIl over 120 years of age (Table 2.1). In the faH of 1999, about one-third ofthe total wood volume was removed from each of the managed sites by the local forest company (P. Wensink, Domtar Inc., 2003, personal communication).

Sample Collection Coleoptera were coIlected using two trap types. A large-area flight-intercept trap (FIT) was based on the design ofPeck and Davies (1980). Two 1.85 m pieces ofrebar were inserted into the forest floor (approximately 2.13 m apart). A 1.25 m x 1.85 m mesh panel of standard black window screening was stretched taut between the upright pieces of rebar. Traps were stabilized by tying nylon cord from the top of each piece of rebar to nearby trees. A wooden trough (1.85 m x 0.5 m x 9 cm) lined with thick c1ear plastic was

25 placed under the mesh panel. A c1ear plastic rain roof (2.36 m x 1.5 m) was stretched over the trap and tied to nearby trees (Fig. 2.2a). A modified version ofthe trunk-window trap (TT) described by Kaila (1993) was also used. A transparent plexiglass window (23 cm x 14 cm) was attached above an inverted 2L pop bottle with the bottom cut away (Fig. 2.2b). The traps were fastened to trees with nylon cord at about 1.3 m above ground. Because only a small portion of the funnel is in direct contact with the tree trunk the traps target flying insects more than those crawling on the trunk. Trunk traps were placed randomly in each plot (within 25 m ofthe FIT), on dead trees with polypore fungus. Iftrees with fungus could not be located, traps were placed on dead trees (or in one case on moribund trees, since dead trees could not be located). Traps were fastened to multiple tree species in a variety of decay classes, since the goal of the study was to assess the beetle diversity of the entire stand and not just of a particular tree species. Two plots (approximately 30 m radius) were established in each forest site, with one FIT and three TTs in each plot (12 FITs and 36 TTs in total). A saturated salt water solution was used for the collecting fluid in all traps. Traps were operated from 29 April 2003 until3 October 2003 (22 weeks). AlI traps were emptied weekly throughout the season using a modified aquarium net (FITs) or a kitchen strainer (TTs) and collecting fluid was replaced as it evaporated.

Coleoptera Identification Beetles were initially stored in 70% ethanol and specimens oftarget families (Table 2.2) were mounted on pins or points and identified to species. Specimens from the primariIy fungivorous families Staphylinidae, Cryptophagidae and Latridiidae were not inc1uded in the study due to Iogistic and taxonomic problems. Specimens from these families, stored in 70% ethanol, are deposited in the Lyman EntomoIogicaI Museum, McGill University, Ste-Anne-de-Bellevue, QC (LEM). In the family Leiodidae, only the subfamily Leiodinae was inc1uded in the study because it is almost exclusively associated with fungus and slime molds. Other leiodid subfamilies are not considered strictly fungivorous and were excluded. Beetles were determined to be fungivorous based on examination of available literature. Specimens were identified to family and genus using

26 standard references (Downie and Arnett 1996; Arnett and Thomas 2001; Arnett et al. 2002). Species identifications were made using keys and descriptions in taxonomie revisions or by collaborating specialists (S.B Peck., Carleton University). Sorne genera (e.g., Agathidium Panzer, Leiodidae) were identified to morphospecies only. Species identifications were verified by consultation with specialists or by comparison to identified specimens in the Canadian Museum of Nature, Aylmer, QC (CMN), the Canadian National Collection oflnsects, Ottawa, ON (CNC) or LEM. Voucher specimens are deposited in LEM and CMN.

Fungus and Coarse Woody Debris Inventory A quantitative inventory of fungal species diversity and abundance at each site was beyond the scope of this study. However, qualitative data were collected on fungi present in each site. Visible fungi within approximately 25 m of each flight-intercept trap were either collected or photographed on two occasions (August and September). Dried specimens and photographs were later identified to species where possible by Scott Redhead (National Mycological Herbarium, Agriculture and Agri-Food Canada) (Appendix 2.1). This list is far from exhaustive because total fungus search time was no more than 25-30 minutes while walking CWD transects and because the fungal inventory was limited mainly to large, visible fungi and thus smaller, more cryptic species were not generalIy inc1uded. In addition, beetles caught in FITs may be capable of flying farther than 25 metres. The line-intercept method was used to measure CWD volumes. At each plot, three 15.24 m transects were established (alI radiating out from the FIT). AlI dead wood over 6 cm in diameter was measured and recorded. Decay c1ass and tree species were recorded for each log where possible. Volume of CWD was calculated according to the 2 equation: V = i2 Ud /8L), where V is volume, d is the diameter of a piece of CWD, and L is the transect length (Bellhouse and Naylor 1996). CWD volume was calculated for each plot (two plots per site) and averages were calculated for each site.

27 Data Analyses Species abundance data from each trap were pooled throughout the season, resulting in 48 samples (12 FIT samples, 36 TT samples). Except for members of the genera Agathidium and Playdema Laporte and Brullé (which were inc1uded in the analyses as morphospecies), genera that could not be identified to species were exc1uded from all analyses. The number of species observed (raw species richness) in each site was recorded. Rarefaction was used to estimate the expected species richness in each site. Rarefaction can also be considered a diversity index because it accounts for both species richness and abundance (Hammond et al. 2004; Olszewski 2004). Standard deviations were plotted for each point on the rarefaction curve. For rarefaction calculations, data from FITs and TTs were pooled by site, resulting in six "samples". Rarefaction was performed using ECOSIM 7.0 (Gotelli and Entsminger 2001). Fixed, individual-based sub-samples (at increments of50) were used in the calculation and the data matrix was randomized 1000 times. To examine evenness and dominance of the data, Fisher's a and Simpson's indices were calculated using EstimateS 6.0b1 (Colwell 2001). Fisher's a is one of the most robust diversity measures and is relatively unaffected by sample size (Magurran 2004). Simpson's index was chosen as a measure of dominance because it performs well at small sample sizes (Magurran 2004). For these calculations, FITs and TTs were treated as individual samples (2 FITs and 6 TTs for each site). ~-diversity was calculated for each pair of sites (with pooled data from both FITs and TTs) using the Morisita-Hom index (CMH). This index was chosen as a similarity measure because it is not strongly influenced by species richness and sample size (Wolda 1981; Magurran 2004). Because the data did not display enough variation (i.e., a low number oftraps collected a large number of species and specimens) an ordination could not be performed. Instead, c1uster analysis was performed on the data from FITs to compare the similarity ofbeetle assemblages between the twelve traps. A Sorenson distance measure and a group averaging linkage method were used in PC-Ord 4.0 (McCune and Mefford 1999) to run the c1uster analysis. Indicator species analysis (Du frêne and Legendre 1997) was also conducted using the program PC-Ord 4.0 to determine ifbeetle species showed significant associations with forest type. Indicator species analysis uses relative

28 abundance and proportional frequency of each species in each sample and produces an indicator value (IndVal) for each species ranging from 0 (no association) to 100 (extremely strong association). An IndVal of 100 means that the presence of a species points to a particular habitat (or group) without error (McCune et al. 2002; Hammond et al. 2004). Rare species (with only a few occurrences) do not yield an IndVal higher than expected by chance (McCune et al. 2002). Indicator values were tested for significance using a Monte Carlo randomization (with 1000 permutations) which resulted in a p-value for each IndVal (McCune et al. 2002; Hammond et al. 2004). Because fungivore catches in TTs were infrequent and patchy, c1uster analysis and indicator values were calculated only for FIT data.

RESULTS A total of Il,888 fungivorous beetles representing Il families and 73 species was collected. Complete species lists for each forest site and the number of specimens in each trap type in each week are in Appendices 2.2a to 2.2f. Traps in 'old-growth' forests collected slightly more individuals (6107 individuals) than those in managed forests (5781 individuals). Flight-intercept traps collected far more individuals (10,896) and species (68) than did trunk-window traps (992 individuals, 55 species). The total number of species did not differ between forest type, with 69 species occurring in both managed and 'old-growth' forest.

Diversity indices for each ofthe six sites are in Table 2.3. Values ofFisher's Ci. at each site ranged from 10.13 to 10.98, the exception being site QG-3, which had a much lower Ci. value of 8.38. The Simpson's index resulted in a range ofvalues from 3.90 to 7.69. Results from both indices ranked QG-3 lower than all other sites. This site had the fewest species (48) but the most individuals (2560), accounting for the low diversity. The low value of the Simpson's index (3.90) reflected the dominance oftwo species, Leiodes subtilicornis Baranowski and Anisotoma horni Wheeler, at this site. Site OG-3 had 70% of the total number ofindividuals of L. subtilicornis. Anisotoma horni was extremely abundant in all sites; however, 25% of all individuals of this species were collected at site QG-3.

29 The mean Fisher's a value for 'old-growth' forest was 9.98 and 10.47 for managed forest. The mean value for the Simpson's index was 5.27 for 'old-growth' forest and 5.76 for managed forest. Thus, managed forests had slightly higher diversity than 'old-growth' forests, although intra-site variation precluded a conclusive result. Rarefaction-estimated species richness for each site (compared at 1550 individuals) is in Table 2.3 and rarefaction curves are in Fig. 2.3a. Standard deviations for each rarefaction estimate are also presented and allow for conclusions as to the significance of each comparison. On a site by site basis, significant differences were as follows: OG-l was significantly higher than MF -1; OG-2 was significantly higher than MF-l and MF-3; MF-3 was significantly higher than MF-l and MF-2; and OG-3 was significantly lower than aIl other sites. A second rarefaction based on pooled managed versus 'old-growth' sites (compared at 5000 individuals) showed that species richness of fungivorous Coleoptera did not differ between forest types (Fig. 2.3b). {3--diversity did not differ between 'old-growth' and managed sites, nor did any pair of sites demonstrate high {3--diversity, with CMH values ranging from 0.852 to 0.969. The cluster analysis (Fig. 2.4) showed no distinction between managed and 'old-growth' sites, with the catch from aIl FITs arising very early from one main branch of the dendrogram. The dendrogram also demonstrated that a few sites show high within-stand variation, as traps from OG-2, OG-3 and MF-3 did not cluster closely together. The indicator species analysis of the fungivorous beetles collected with FITs is in Table 2.4. Only species with p-values ::;;0.5 are included in the table. Five species showed a significant association with either managed or 'old-growth' forests (p ::;;0.05). One species, Triplax macra LeConte, was very strongly associated with 'old-growth' forests. Four species showed good to moderate associations with managed forests: Agathidium sp. 1, Anisotoma blanchardi (Hom), Anogdus obsoletus (Melsheimer), and Mycetina perpulchra (Newman). Members of the genus Anisotoma were collected from May/early June until July/early August with relative abundance peaking in mid to late July. Some species of Agathidium, however, did show phenological differences (e.g., Agathidium pulcrum LeConte was not collected after July, while Agathidium sp. 3 was not collected before July). Colenis impunctata LeConte and Hydnobius longidens LeConte were collected

30 throughout the sampling season, from May to September/October. Species of Anogdus, Liocyrtusa and Lionothus were collected from mid-June until September/October. Leiodes pygmaea (Baranowski) and Leiodes sorenssoni (Baranowski) were collected from mid July/early August until October, while Leiodes subtilicornis was found from early June until October, with numbers being highest in late June and mid September. Mycetina perpulchra, Tritoma pulchra Say, Mycetophagus jlexuous Say, Scaphidium quadriguttatum Melsheimer and Tritoma mimetica (Say) were collected throughout the season, from May to October. Triplax macra was taken from mid May untillate August, its numbers peaking in early June and mid August - these peaks may correlate with fruiting times of its host fungus. Volume of coarse woody debris was much higher in 'old-growth' sites than in managed forests (190.32 m2 ± 85.42 for 'old-growth', 63.21 m2 ± 28.79 for managed). This was mostly due to 'old-growth' forests having more downed 10gs in large diameter classes. A1though managed forests had a few large downed 10gs, the majority of wood encountered on the transects was fine woody debris (i.e., 10gging slash).

DISCUSSION Comparison of Forest Types

Fisher's Cl, Berger- Parker and rarefaction dea1 with a-diversity, the diversity of species within a community or habitat (Southwood 1978). Rarefaction, whi1e providing un-biased estimates of species richness at each site, a1so allows for standard deviations to be ca1cu1ated for each estimate, faci1itating a comparison between a-diversity in each forest site and between each habitat. In contrast, {3-diversity is a measure of the rate and extent of change in species a10ng a gradient from one habitat to another (Southwood 1978). Distributions ofpairwise {3-diversity measures (such as those generated by the Morisita-Hom index) can be compared directly (Magurran 2004). C1uster ana1ysis is another intuitive, simple method to compare {3-diversity by graphically representing differences amongst samples/sites (Magurran 2004). Sites or samp1es that c1uster together are more similar to one another than to those that are located at a distance from each other.

31 In this study, an methods used to analyse the data - diversity indices (Table 2.3), rarefaction (Fig. 2.3), {j-diversity and cIuster analysis (Fig. 2.4) - indicated that there was no significant difference in the species richness and relative abundance of fungivorous Coleoptera between managed and 'old-growth' sites. This was contrary to my initial prediction. One explanation may be that the families ofbeetles examined in this study are not vulnerable to the forest management practices carried out in these study sites. Second, the three managed sites are generally managed for 'old-growth characteristics', meaning that a certain percentage of dead wood (snags and CWD) is maintained in the stands as wildlife habitat. This 'leftover' wood may also provide sufficient habitat for the fungivorous beetles (and their host fungi) examined in this study. Third, the effect ofthe cutting may not yet have had time to influence the beetle communities in the managed forests, or any negative effects that forest harvest had on the beetle community may have already been mitigated. Fourth, it is possible that specialist beetle species within these families that may have been vulnerable to a difference between forest types, were extirpated from all six sites prior to this study, due to the degree of fragmentation and isolation that has occurred. Finally, it is possible that the field component ofthis study was conducted in an atypical year and that, had additional seasons been sampled, different diversity patterns may have been detected. The effect of fragmentation on arthropod populations has been widely studied and many believe that the survival ofrare and/or specialist species is determined by metapopulation dynamics (demography of isolated/fragmented populations) (Siitonen and Martikainen 1994; 0kland et al. 1996). All sites were very small (between 5.9 ha and 13.8 ha) forests, had little interior habitat and a lot of edge; this may have had a strong effect on beetle species composition (Murcia 1995; Niemela 1997; Jokimaki et al. 1998). However, the effect of edge on forest species is still controversial and leiodids in particular are fairlyactive fliers (Chandler and Peck 1992) and thus dispersal may not be a large factor determining the species composition of these beetles in forest fragments. There was, however, significant inter- and intra- site variation between and within the 'old-growth' sites and this may have resulted in inconcIusive results for the (X diversity between managed and 'old-growth forests. Diversity indices ranked site OG-3 as lower than all other sites (whether 'old-growth' or managed). There are several

32 possible reasons for this difference. First, the site, although having the same basic tree species composition and age as the other 'old-growth' sites, has a much more open canopy, allowing light to penetrate through to the ground. Second, unlike other 'old growth' sites there was very little regeneration ofbeech and hemlock at this site and thus, the understory was relatively open. Third, the site remained moist much later into the season than did the other sites. This site may be undergoing a forest-to-wetland successional process, and this likely affected the beetle species in the site. The dominant species at the site support this contention. Over 70% of individuals of Leiodes subtilicornis collected were caught at site OG-3. Adults and larvae of Leiodes spp. are known to feed on hypogean and subterranean fungi (Newton 1984; Baranowski 1993). Most species of Leiodes are thought to prefer open areas (Chandler and Peck 1992; Baranowski 1993) and this potentially explains their abundance at this site. The cluster analysis also indicated intra-site variation. The two FITs in MF-l, MF-2 and OG-l each clustered together on the dendrogram (Fig. 2.4); traps in sites MF- 3, OG-2, and OG-3, however, did not cluster together. This illustrates the importance of sampling over a wide area if the goal is to characterize a particular habitat. Many habitats are composed of a variety of different microhabitats. If only one FIT trap had been placed in each forest site an incomplete picture of the species composition and distribution at the sites may have resulted.

Species Diversity of Leiodinae The round fungus beetles (Leiodidae) were the most abundant family collected, accounting for 87.4% of all specimens identified. Only members of the subfamily Leiodinae, containing many species that are specialists on slime molds (Wheeler 1984), were considered; the other subfamilies (e.g., Cholevinae) do not feed exclusivelyon fungus or slime molds (Peck and Cook 2002). It has been hypothesized that slime molds, which are associated with downed woody debris (Stephenson 1988), may be more abundant in 'old-growth' forests due to the higher volumes ofCWD available for their development (Chandler and Peck 1992). It is thus logical to hypothesize that slime mold specialists may also be more diverse and abundant in 'old growth' forests. In this study, however, strong associations between slime mold-feeding

33 beetles (e.g. Agathidiurn, Anisotorna, Sphindidae) and 'old-growth' forests were not evident. In fact, two species (Agathidiurn sp.l and Anisotorna blanchardi) showed significant associations with managed forests (Table 2.4). While sorne species did show greater abundance in 'old-growth' forests (e.g., Agathidium sp. 4, Anisotorna discolor (Melsheimer), Anisotorna inops W.J.Brown) the same was also true for species in managed forests (e.g., Agathidiurn sp.l). Anisotorna inops is a species considered a potential 'indicator' of old-growth forests in north-eastem North America (Chandler and Peck 1992). Anisotorna inops did not show a strong association with 'old-growth' forests in this study; however, this was probably because stands were approximately the same age (Chandler and Peck (1992) compared old and young forests) andA. inops maybe associated with mature forests, managed or not. It is believed that many slime molds favour a particular substrate; however, little is known about the specificity of slime mold species to any one particular substrate (Blackwell1984). It is possible that slime molds can develop just as efficiently on FWD (which is common in managed forests) as they do on CWD (which is usually more common in 'old-growth' forests). To our knowledge, there is no published literature exploring the size classes of dead wood utilized by slime molds. This may explain why there was no apparent difference in the species richness and abundance of slime mold feeding beetles between 'old-growth' and managed forests. Only one leiodid species,

Anogdus obsoletus, had a significant association with managed forests (IndVal = 77.9, P = 0.011). Unfortunately, there is little literature describing fungal associations and feeding strategies ofthis genus/species and it is difficult to speculate on why the species was associated with managed forests.

Species Diversity of Other Fungivores Other families cOllected in high numbers included the Erotylidae (pleasing fungus beetles), Mycetophagidae (hairy fungus beetles) and Endomychidae (handsome fungus beetles) (Table 2.2). AIl species collected from these families are mycophagous to sorne degree on a variety offungal species (e.g., Lawrence and Newton 1980; Newton 1984; Skelley et al. 1991). For sorne species the beetle-fungus association is weIl known (e.g., Lycoperdinaferruginea LeConte on Lycoperdon pyriforme Schaeff), while for many

34 others, virtually nothing is known (e.g., Danae testacea (Ziegler)). In general, the Endomychidae and Mycetophagidae were more abundant in managed forests and the Erotylidae were more abundant in 'old-growth' forests. Onlyone species, Mycetina perpulchra, showed a significantly strong association with managed forests (IndVal =

78.8, P = 0.013). This species is widely distributed and is presumed to be at least partially polyphagous on a wide range of fungi (F. Shockley 2004, personal communication). In contrast, Triplax macra was strongly associated with 'old-growth' forests

(IndVal = 91.4, p = 0.017). The genus Triplax is divided into two groups: the macra group (associated with the polypore genus [nonotus P.Karst) and the thoracica group (associated mainly with gilled Pleurotus Fries fungus) (Skelley et al. 1991; Goodrich and Skelley 1993). Anecdotal evidence indicates that [nonotus spp. may be more readily encountered in old-growth forests (F. Shockley 2004, personal communication), and this may explain the strong association of T. macra with 'old-growth' forests. No species of [nonotus were identified during this study, although the inventory was not exhaustive. Pleurotus spp., however, were frequently encountered fruiting on dead standing beech trees, which were common in 'old-growth' forests and a much stronger component of 'old-growth' forests than ofmanaged sites (Table 2.1). Deciduous trees have long been recognized as being important sources of dead wood for saproxylic insects (Siitonen and Martikainen 1994; 0kland et al. 1996). It is like1y that dead beech serves an important function in these forests in terms of fungus substrate and beetle habitat. Thus, the association of T. macra with 'old-growth' forests may be coincidental, the difference being more related to the high quality ofbeech than to any harvesting effect. The fact that T. macra was not collected in sites MF-1 and MG-2 (Table 2.2), sites which did not contain a high percentage ofbeech, supports this possibility.

Trapping Differences Although beetles caught in flight-intercept traps accounted for over 90% of the individuals collected, several species were collected more frequently in TTs than in FITs. (e.g., Anisotoma inops and Triplax thoracica) (Table 2.2). Euparius marmoreus (Olivier) (Anthribidae) and Agathidium sp.1 were caught exc1usively in TTs. Species from the

35 families Tenebrionidae and Trogossitidae were rarely collected but were caught more frequently in TTs. This was also the case for Ranidea unic%r (Endomychidae), Dacne quadrimaculata (Say) (Erotylidae), and Anisotoma g/obososa Hatch (Leiodidae). The leiodid subterranean and hypogean fungus feeders, e.g., Leiodes, Cyrtusa, Co/enis, and Anogdus, were never collected with TTs. There were marked differences in the abundance of Anisotoma species collected with TTs versus FITs: 1) Anisotoma disc%r (Melsheimer) and Anisotoma basa/is (LeConte), while abundant in both traps, were more frequently collected with FITs; 2) Anisotoma horni, the most abundant species caught throughout the study, was collected almost exc1usively in FITs; and 3) Anisotoma inops was collected far more often in TTs. Approximately 30% ofthe species collected were represented by fewer than ten individuals and over half ofthese species were represented by less than five individuals. Flight-intercept traps are among the best traps for collecting large numbers of fungivorous beetles (Peck and Davies 1980); however, due to their large size, they also may collect many transient species. One ofthe problems associated with trap data is that many species are wide-ranging 'tourists' and can sometimes be found in habitats where they do not reproduce or develop (NiemeUi 1997; Martikainen et al. 1998). A difficulty arises when trying to decide which species are 'tourists' and which species are simply rare and thus uncommonly collected. Often it is difficult to make such conclusions in the absence of additional sampling data, especially if little is known of the biology of each species. For the species collected during this study, members ofthe Tenebrionidae and Anthribidae were infrequently collected using FITs and are likely not nearly as abundant as the Leiodidae, Endomychidae, Erotylidae and Mycetophagidae. However, several species within these abundant families (e.g., the endomychid Ranidea unic%r (Ziegler) and the leiodids Leiodes campbelli Baranowski and Triarthron pennsy/vanicum Hom) were represented by very low numbers and, since host and habitat records are not always known, these species may weIl be transient individuals.

Conclusions: Fungus-insect Relationships and Forest Management This study did not provide an in-depth inventory of fungi present at each site. Like arthropods, fungi are extremely diverse but they are inadequately studied and large

36 gaps remain in the taxonomy and natural history ofmany 'species'. There are several reasons for this infonnation gap: many fungi are smaIl and inconspicuous and are easily overlooked; many fungi produce ephemeral fruiting bodies and thus require complete seasonal sampling; species concepts are difficult to apply; and systematic mycology is underfunded (Cannon 1997). Given that many of the same problems plague arthropod and fungal research, it is not surprising that large gaps remain in our understanding of fungus-insect relationships. In the future, it would be valuable to expand studies such as this one to inc1ude an extensive fungal inventory as weIl as rearing experiments. Without such research, our understanding of the complex fungus-beetle relationships in forests will be incomplete. Forest management did not have a significant effect on the species richness and relative abundance of fungivorous beetles in this study. However, there were differences in the species composition ofbeetles between forest types (e.g., Triplax macra was more common in 'old-growth' sites and Mycetina perpulchra in managed sites). These results are in accordance with similar studies that have occurred in the GLSL region - e.g., ground-occurring beetles (Werner and Raffa 2000) and leiodid beetles (Chandler and Peck 1992). Studies which have found significant differences in Coleopteran diversity commonly look at the effect of extreme forest management practices such as c1earcuts (e.g., Kaila et al. 1997; Niemelli et al. 1993), or at the differences between stands of distinctly different ages (e.g., Martikainen et al. 2000). In this study stands were approximately the same age and forest management was the only variable. It is possible that a greater stand age difference would have produced significant differences in species diversity. 'Old-growth' forests did have higher volumes of coarse woody debris than did managed forests, but there was great variation in these values from plot to plot. Further study is required in order to detennine if the larger volume of dead wood in these forests is linked to higher diversity and abundance offungus. Many forest systems are managed to maintain 'old-growth characteristics' - i.e., a set volume or number of dead or dying trees (snags, downed wood, cavity trees) are retained in the harvested forest stands. These structures provide important habitat for a variety ofwildlife species, inc1uding saproxylic beetles (0kland et al. 1996; Simila et al. 2003). In populated areas like southern and south-eastern Ontario, where tracts of forests

37 are small and fragrnented, these habitat characteristics become ever more important. Data from this study suggests that managed forests, being managed to retain important habitat structures, are also helping to maintain biodiversity. It is especially important to manage forests with the goal of maximizing suitable habitat and biodiversity in the GLSL region of south-eastem Ontario. Forests is this area are small and fragrnented (as opposed to forests in the boreal zone which tend to be larger more continuous tracts) and this makes it much more difficult for species to recolonize sites after local extinctions occur.

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44 Table 2.1. Location, tree species, size, age and general description of study sites.

Site Location SEecies CornEosition* Size A~e General Information OG-l Cornwall, ON He37Mr3SBel7Pw90h2 9.3 ha 160-210 Private ownership. Stand Iikely 45°02.l60'N years selectively harvested about 160 years 74°47.470'W ago (high-grade - took only largest and best trees). Surrounded by sugar rnaple forest, city roads, and agriculturalland. Acidic soil, fresh to rnoist; u)2lands sand~ loam. OG-2 Lancaster, ON He3SBe3SMhlsPwloOhs -12 ha + 120 Owned by St. Lawrence Parks 45°07.750'N years Commission ("Raisin River Park", 74°30.875'W (climax operations closed 1992). Sorne forest) evidence of thinning in past. Surrounded rnostly by Hwy 401 and the Raisin River. Flat, low-lying terrain. OG-3 Morrisburg, ON He3oBe30Mh300hlO - 8ha + 130 Owned by St. Lawrence Parks 44°56.750'N years (Jate Commission ("Upper Canada Sugar 75°06.035'W succession Bush"/"Crysler Farm Battlefield Park al forest) Forest"). On west edge of80 ha sugarbush; county roads and clearings to east, north and south of site. Mesic to wet; hummocky sandy loarn over cla~ )2lain. MF-l Lancaster, ON He4oMh2oAwloMrlOPw 13.80 + 100 Private ownership. Managed by 45°1O'N lOOh lO (Be, Ob, By, ha years Dorntar Inc. Woodlot was selectively 74°27'W Bd, Bn, Bf, Cw) cut in 1988. Surrounded by agriculturalland (corn, soy). Sandy loam; sIightl~ acidic soil. MF-2 Bainsville, ON He4oMr3oMh200hlo(A 11.30 150 years Private ownership. Managed by 45°12.01O'N w, Bd, Bf, Be, By, Cb, ha Domtar Inc. Stand was harvested 74°25.650'W Brn, Po) -40yrs ago to build barns on farm. Surrounded by agriculturalland (corn, soy). Very fine sandy loarn; sorne silt cla~. MF-3 Bouck's Hill, He3oMh3oBd2oBeloIdio 5.9 ha 150 years Private ownership. Managed by ON (Hb, Aw) Dorntar Inc. Stand was once part of 45°00'N large maple syrup operation. 75°12'W Surrounded by agricultural lands {ha~, so~2. Cla~ loam.

* Species codes: He = eastem hemlock; Be = American beech; Mh = hard (sugar) maple; Mr = red maple; Pw = white pine; Aw = white ash; By = yellow birch; Bd = basswood;

Ob = bur oak, Bn = black oak; Rb = bittemut hickory; Bf = balsarn tir; Cw = white cedar;

Cb = black cherry; Em = elm; Po = poplar; Id = ironwood; Oh = other hardwood. Subscript nurnbers following species code indicate percent oftotal tree species composition occupied by that species in the stand.

45 Table 2.2. Fungivorous Coleoptera collected at study sites, 2003. Specimens in trunk window traps are in parentheses; those in flight-intercept traps are not. Totals for each species and each site are pooled numbers from both trap types. Species in bold were primarilyor exclusively collected in trunk window traps. Numbers in brackets following family names indicate the total number of individuals and the total number of species collected from each family.

Site S~ecies MF-1 MF-2 MF-3 Total OG-1 OG-2 OG-3 Total Family Anthribidae (47, 5) Choragus sayi LeConte (1 ) 1 1 1 Euparius marmoreus (Olivier) (2) 2 (3) (1) (3) 7 Euxenus punctatus LeConte 1 Tropideres dorsalis (Thunberg) 1 Troeideres tricarinetus (Pierce} 9 1 11 7 3 13 23 Family Cerylonidae (89, 3) Cerylon castaneum Say 5 3 (1) (1 ) 10 (2) 2 Cerylon unicolor (Ziegler) 10 (2) 7 (1) 2 (4) 26 15 (2) 8 (5) 11 (1) 40 Philothermus g.1ebriculus LeConte 4 1 5 3 1 4 Family Endomychidae (272, 6) Danee testacee (Ziegler) 1 1 15 15 Endomychus b/guttatus Say 1 (1) 5 (3) 14 (1) 25 2 (2) 1 (2) 7 Lycoperdina ferruginea LeConte 7 9 1 17 1 2 3 Mycetina perpulchra (Newman) 74 (1) 30 45 (1) 151 18 13 10 41 Phymaphora pulchella Newman 2 (1) 3 6 2 3 (1) 6 Ranidea un/color ~1!!Iler} {1 } 1 Family Erotylidae (569, 8) Dacne quadrimacu/ata (Say) 2 (1 ) 3 1 1 (8) 2 (1) 13 Triplax dissimulator (Crotch) 1 1 Triplax flavicollis Lacordaire (2) 2 2 6 10(11 ) 1 22 Triplax frosti Casey 1 1 1 (2) 1 4 Triplax macra LeConte 7 7 45 (1) 8 (4) 21 (1) 80 Triplax thoracica Say 1 (14) 1 5 21 17 (81) (1 ) 3 102 Tritoma mimetica (Say) 4 2 26 32 19 10 16 45 Tritoma F!!!./chra Sa~ 16 {2} 24 42 (1} 85 43 35 {1} 66 {2} 147 Family Leiodidae (Leiodinae) (10386, 38) Agathidium sp.1 76 111 113 300 82 26 20 128 Agathidium sp.2 67 81 44 192 51 42 21 114 Agathidium sp.3 18 12 2 32 18 16 18 52 Agathidium spA 4 (7) 5 (1) 11 (1) 29 86 (5) 10 (3) 12 (2) 118 Agathidlum sp.5 15 (1) 25 18 59 56 (3) 14 14 87 Agathidium sp.6 2 3 1 6 2 7 3 12 Agathidlum sp.7 4 (1) 9 1 15 11 1 2 14 Agathidium sp.8 3 1 2 6 3 3 Agathidium sp.9 29 13 9 51 4 3 10 17 Agathidium sp.10 1 1 2 2 1 (1) 4 Agathldium sp.11 (2) (2) 4 (1 ) (1 ) (2) 4 Agathidium sp.12 3 9 1 13 1 1 Agathidium depressum Fall 4 1 3 8 2 (1) 5 (1) 9 Agathidium pulchrum LeConte 3 6 11 20 17 1 25 43 Agathidlum ?oniscoides (1 ) 1 1 (1) 4 1 3 (2) 5 11 Ag/yptinus /aevis (LeConte) 1 1 Anisotorna basa/is (LeConte) 36 (48) 82 (55) 50 (7) 278 80 (41) 20 (38) 58 (10) 247 Anisotoma blanchardi (Hom) 76 135 61 272 45 45 59 149 Anisotoma discolor (Melsheimer) 151 (27) 51 (14) 118(16) 377 139 (5) 231 (64) 211 (10) 660 Anisotoma geminata (Hom) 9 2 2 13 4 1 5 Anisotoma globososa Hatch 1 1 1 (5) 8 2 (1 ) 3 Anisotoma homiWheeler 574 (33) 799 (10) 828 (9) 2253 628 (3) 679 (9) 1206 2525 Anlsotoma inops W.J.Brown 3 (20) 2 (5) 42 (64) 136 33(115) 30 (44) 21 (28) 271 Anogdus puritanus (Fall) 28 94 31 153 12 27 1 40 Anogdus obsotetus (Melsheimer) 60 22 34 116 3 15 15 33 Co/enis impunctata LeConte 40 116 121 277 127 15 68 210

46 Table 2.2. continued

Seecies MF-1 MF-2 MF-3 Total OG-1 OG-2 OG-3 Total Family Leiodidae (Lelodinae) (continued) Cyrlusa sub/es/acea (Gyllenhal) 2 3 2 1 4 7 Hydnobius n. sp. 2 10 12 1 1 Hydnobius longidens LeConte 11 27 38 2 2 4 Isoplas/us (ossor Hom 1 4 46 51 1 1 Leiodes campbelli (Baranowski) 2 1 3 3 3 Leiodes pygmaea (Baranowski) 4 4 11 19 73 73 Leiodes sorenssoni (Baranowski) 11 3 14 34 34 Leiodes subtilicomis (Baranowski) 45 36 49 130 13 17 381 411 Liocyrtusa nigriclavis (Hlisnikovsky) 10 66 76 30 30 Liono/hus (orlicomis Daffner 4 2 56 62 16 6 2 24 Triarlhron I2!!.nnsylvanicum Hom 1 1 3 1 4 Family Mycetophagidae (391,4) Myce/ophagus flexuosus Say 88 (9) 70 (17) 35 (4) 223 10 (5) 26 (11) 48 (2) 102 Myce%phagus pluripunctatus (1 ) 1 2 (1 ) 1 2 LeConle Mycetophagus punctatus Say 9 13 (2) 10 (3) 37 3 (3) 1 (4) 12 23 Mypetoehag.us serrulatus Case): 1 1 1 1 Family Scaphidiidae (73, 1) Scaphidium quadriguttatum 15 14 14 43 15 7 8 30 Melsheimer Family Sphindidae (33, 1) Odontosphindus denticollis 3 (4) (1 ) 6 (6) 20 2 4 (1) 6 13 LeConle Family Tenebrlonidae (19, 5) Bolitoph.gus corticol. Illiger (1) 1 Bolitophagus comutus (Panzer) (3) 3 (1) 1 Diaperis macu/ata Olivier (1) 1 (2) 2 (1 ) 5 Neomid. bicomis (Fabriclus) (2) 1 (1 ) 4 PI.tydema seA {1 } 1 (1 } 1{1} 3 Family Trogossitidae (8, 2) Tenebroides ?corticalis (1 ) 1 (3) (1 ) 4 (Melshelmer) Thymalus marg/nicollis (1 ) 2 (1 ) 1 Chevrolat Family Zopheridae (1,1) Phelloesis obcordata (Kirb):l 1 1 TOTALS 1702 1959 2120 5781 1983 1564 2560 6107

47 Table 2.3. Raw species richness (Sobs), number ofindividuals, rarefaction estimated species richness (species ± sn, standardized to 1550 individuals), and diversity indices of fungivorous Coleoptera at study sites (both trap types pooled).

Site Sobs Numberof Rarefaction Fisher's a Simpson's individuals estimated species collected richness OG-l 56 1983 53.35 {± 1.11} 10.71 7.69 OG-2 54 1564 53.84 {± 0.08} 10.84 4.22 OG-3 48 2560 44.05 {± 1.42} 8.38 3.90 MF-l 52 1702 51.10 {± 0.40} 10.13 6.45 MF-2 57 1959 53.08 {± 1.64} 10.98 5.12 MF-3 55 2120 51.35 {± 1.47} 10.31 5.70

48 Table 2.4. Indicator species analysis of fungivorous Coleoptera collected in flight

intercept traps. Species in bold have a significant (p ~.05) Indicator Value (IndVal); species with p > 0.5 are not shown.

Forest Family Species Total No. Indicator p- T~~e collected value value 'Old-growth' Leiodidae Agathidium sp.3 61.9 0.296 (Leiodinae) Agathidium sp.4 84.4 0.104 Agathidium sp.6 44.4 0.365 Agathldium ?oniseoides 54.5 0.285 Agathidium pulchrum LeConte 56.9 0.451 Anisotoma discolor (Melsheimer) 64.5 0.084 Anisotoma inops W.J.Brown 64.1 0.356 Erotylidae Trlplax maera LeConte 91.4 0.017 Tritoma pulchra Say 63.7 0.086 Anthribidae Tropideres tricarinatus (Pierce) 67.6 0.152 Cerylonidae Cerylon unic%r (Ziegler) 64.2 0.171 Tenebrionidae Diae.eris macu/ata Olivier 33.3 0.451 Managed Leiodidae Agathidium sp.l 70.1 0.050 (Leiodinae) Agathidium sp.2 62.7 0.069 Agathldlum sp.8 44.4 0.240 Agathidium sp.9 75.0 0.089 Agathidium sp.12 61.9 0.142 Anlsotoma blanehardl (Horn) 64.6 0.044 Anogdus obso/etus (Melshelmer) 77.9 0.011 Anogdus puritanus (Fall) 79.3 0.074 Hydnobius longidens LeConte 60.3 0.191 Hydnobius n. sp. 46.2 0.182 Isop/astus fossor Horn 65.4 0.109 Lionothus nigric/avis (Hlisnikovsky) 47.8 0.313 Endomychidae Endomychus biguttatus Say 72.5 0.144 Lycoperdina ferruginea LeConte 70.8 0.081 Myeetlna perpulehra (Newman) 78.8 0.013 Mycetophagidae Mycetophagus flexuosus Say 69.7 0.053 Mycetophagus punctatus Say 66.7 0.160 Ce!i:lonidae Cet;l}on castaneum Sa;l 33.3 0.453

49 --

QarI.- OfY__ _ """'---, F.-oIV_ 'Old-growth' l.OoIMtII tO 0 10 :zo _~I Forest • -<",'C',

Figure 2.1. Location of study sites, eastem Ontario, Canada

50 A

Figure 2.2. A - Flight-intercept trap (FIT); B - Trunk-window trap (TT).

51 60 ~------,

III III cCI) .c 40 Co) Ëi III CI) 'u 30 c.CI) fi) --.-MF-2 i ___ MF-3 Co) 20 -CI) c. -e-MF-1 w>< -6-0G-1 --er-OG-2 10 ---e-- OG-3

o +------_.------~------_.------~------~------~ o 500 1000 1500 2000 2500 3000 Num ber of Individuals

Figure 2.3a. Rarefaction estimates of expected species richness (± 1 SD) of fungivorous Coleoptera collected with flight-intercept and trunk-window traps in six forest sites.

52 80

CIl CIl 60 CI) C oC u i2 50 CIl CI) 'C:; CI) 40 Co t.n "S 30 u a.CI) ~ 20 ___ M anaged forest 10 -e- 'Old-growth' forest

0 0 1000 2000 3000 4000 5000 6000 Num ber of Individuals

Figure 2.3b, Rarefaction estimates of expected species richness (± 1 SD) of fungivorous Coleoptera collected with flight-intercept and trunk-window traps in managed versus 'old-growth' forests.

53 Percent Similarity

100 75 50 25 o

MF-1a 1 1 MF-1b 1 MF-2a 1 MF-2b OG-1a OG-1b 1 OG-3b 1 MF-3a OG-2a MF-3b OG-2b OG-3a

Figure 2.4. Dendrogram of c1uster analysis on species of fungivorous Coleoptera collected from flight-intercept traps.

54 Appendix 2.1. Fungi found within an approximate 25m radius ofthe flight-intercept traps at each site.

Site Species/Genera

OG-l ?Arcyria sp., Climacodon septentrionale, Fomesfomentarius, Ganoderma applanatum, Hygrocybe sp., Hypocrea sp., Hypoxylon sp., Laetiporus sp., Lycogota epidendron, Phe/linus sp., Pleurotus sp., Russula sp., Trametes sp., Trametes versicolor, Trichaptum sp., Trichaptum biennis, ?Tyromyces sp.

OG-2 Fomesfomentarius, Hygrocybe spp., Mycena sp., Oxyporus sp., ?Peziza sp., Russula sp., Tetrapyrgos sp., ?Tylopilus sp.

OG-3 ?Clitocybula sp., Fuligo septica, Ganoderma applanatum, Ganoderma tsugae, Hericium sp., Hygrocybe sp., Hypoxylon sp., Lactarius sp., Phaeolus schweinitzii, Phe/linus sp., Pluteus sp., Russula sp.,

MF-l Amanita ?flavoconia, Ceratiomyxa ?fructiculosa, Crinipe/lis ?zonata, ?Daldina sp., Exidia sp., Fomesfomentarius, Ganoderma sp., ?Lenzites sp., Marasmius rotula, Marasmius ?siccus, Mycena sp., Mycena haematopus, Mycena leaiana, Phe/linus sp., Physalacria inflata, Piptoporus betulinus, Pluteus sp., Schizophyllum commune, Scute/linia sp., Stemonitis sp., ?Trametes sp., Trichaptum sp., Xerocomus sp., Xerula sp.

MF-2 Clitocybe sp., Coprinus sp., Crinipe/lis zonata, Ganoderma sp., ? Lepiota sp., Lycoperdon sp., Marasmius sp., Mycena leaiana, Pluteus sp., Russula sp., Sc/eroderms sp., Tetrapyrgos sp., Terapyrgos nigripes, Trametes versicolor

MF-3 Amanita sp., Ganoderma sp., Gloeophyllum sp., Hydnellum sp., Hygrocybe sp., Peziza sp., Russula sp., Tremella con cres cens

55 Appendix 2.2a. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #1, Cornwall, ON, (OG-l).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S~ecies May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sel! Sel! Sel! Sel! Oct Family Anthribidae Choragussay; LeConte Eupariusmarmoreus (Olivier) (1 ) (1 ) (1) Euxenuspunctatus LeConte Tropideresdorsalis (Thunberg)

Tropiderestricarinatus (Pierce) 2 2 Family Cerylonidae Ceryloncastaneum Say Cery/onunico/or (Ziegler) (1 ) 6 5(1) 3 Philothermusglabriculus LeConte VI 2 0\ Family Endomychidae Danaetestacea (Ziegler) Endomychusbiguttatus Say (1) (1) 1 Lycoperdinaferruginea LeConte Mycetinaperpu/cra (Newman) 5 1 4 5 1 Phymaphorapu/chella Newman Ranideaunicolor (Ziegler) (1) Family Erotylidae Dacnequadrimaculata (Say) Triplaxdissimulator (Crotch) Triplax flavicollis Lacordaire (2) 1(7) 5(2) 2 Triplax frostj Casey (2) Triplaxmacra LeConte 2 2(1) 7 4 5 2 7 2 8 5 Triplax thoracica Say 2(2) 2 2(16) 2(7) 1(4) 1(5) (19) 2(18) (10) 2 1 Tritomamimetica (Say) 1 2 5 2 3 Tritomapulchra Say 3 2 4 3 2 4 5 5 5 5 2 Appendix 2.2a. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S~ecies Ma~ Ma~ Ma~ Jun Ma~ Jun Jun Jun Jul Jul Jul Jul Jul AU9 AU9 AU9 AU9 Se~ Se~Se~ Se~Oct Family Leiodidae Agathidiumsp.1 3 ~ 18 17 8 2 3 6 Agathidiumsp.2 2 8 11 10 4 2 1 2 Agathidiumsp.3 2 3 4 2 2 2 AgathidiumspA (2) 5(1) 12 4 6 6 (1 ) 23(1) 20 2 2 1 Agathidiumsp.5 3 4 12(1) 14 4 2(1) 1 3(1) 6 2 2 1 Agathidiumsp.6 Agathidiumsp.7 6 3 Agathidiumsp.8 Agathidiumsp.9 1 Agathidiumsp.10

Agathidiumsp.11 (1) Agathidiumsp.12 VI -J Agathidiumdepressum Fall Agathidiumpu/chrum LeConte 6 2 6 Agathidium?oniscoides

Ag/ypilnus/aevis (LeConte) 1 Anisotomabasa/is (LeConte) 7(3) 3(2) 3(6) 6(3) 10(1) 6(2) 1(3) 9(5) 9(2) 6(1) (5) 5(4) 4(3) (1 ) Anisotomab/anchardi (Hom) 2 6 5 14 6 5 2 2 Anisotomadisc%r (Melsheimer) 14 7 23(2) 8(1) 9 23(2) 18 3 4 2 3 2 Anisotomageminata (Hom) 1 Anisotomaglobososa Hatch 2 Anisotomahomi Wheeler 7 14 38 75 47 34 37 53(1) 93 (2) 40 46 24 10 4 2 Anisotornainops W.J.Brown 3(8) 1(3) 1(11) 7(7) 2(6) 7(8) (4) 2(9) 1(12) 1(6) (5) 6(18) 1(9) (6) 1(2) (1) Anogduspuri/anus (Fall) 4 5 Anogdusobsoletus (Melsheimer) Colenisimpunctata LeConte 5 3 8 8 12 9 19 8 10 7 3 4 2 3 Cyrtusasubtes/acea (Gyllenhal) Appendix 2.2a. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Leiodidae (continued) Hydnobiusn. sp. Hydnobiuslongidens LeConte Isoplastustossor Hom Leiodescampbelli (Baranowski) 2 Leiodespygmaea (Baranowski) Leiodessorenssoni (Baranowski) 2 9 1 6 2 5 7 LeiocJessublilicomis (Baranowski) 2 2 Liocyrlusanigriclavis (Hlisnikovsky)

Lionothustorlicomis Daffner 2 2 5 Triarthronpennsylvanicum Hom 2 Family Mycetophagidae Mycetophagusflexuosus Say 1(1) (1 ) (1) 2 1 2 2(1) 1(1) VI Myceotophaguspluripunctatus LeConte 00 (1 ) Mycetophaguspunctatus Say (1 ) (1) (1) Mycetophagusserrulatus Casey

Family Scaphidiidae Scaphidiumquadriguttatum Melsheimer 2 1 1 1 4 2 Family Sphindidae Odontosphindusden/icollis LeConte 2 Family Tenebrionidae Bolitophaguscorlico/a IIliger Bolitophaguscomutus (Panzer) (1) Diaperismaculata Olivier (1 ) (1 ) Neomidabicomis (Fabricius) (1) (1) Platydemasp.A (1 ) Upisceramboides Fabricius Appendix 2.2a. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Trogossitidae Tenebroides?corticalis (Melsheimer) (2) (1 ) Thymalusmarginicollis Chevrolat (1 ) Family Zopheridae Phellopsisobcordala (Kirby)

Ut \0 Appendix 2.2b. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #2, Lancaster, ON, (OG-2).

Eupariusmarmoreus (Olivier) (1 ) Euxenuspunctatus LeConte

Tropideresdorsalis (Thunberg) Tropideres tricarinatus (Pierce)

Family Cerylonidae CelY/oncastaneum Say (1 ) (1 ) CelY/onunic%r (Ziegler) (1 ) (1 ) 5 1(1) 1(1) (1) 1 Philothermusg/abricu/us LeConte 0\ o Family Endomychidae Danaetestacea (Ziegler) 3 1 1 4 2 Endomychusbiguttatus Say 1(1) (1) Lycoperdinaferruginea LeConte Mycetinaperpu/cra (Newman) 2 2 2 2 2 Phymaphorapu/chelia Newman Ranideaunicolor (Ziegler)

Family Erotylidae Dacnequadrimaculata (Say) 1(6) (2) Triplaxdissimulator (Crotch) Trip/ax flavicollis Lacordaire Triplax frosti Casey Trip/axmacra LeConte 2(2) 4(2) 2 Trip/axthoracica Say (1) Tri/omamimetica (Say) 1 2 3 Tri/omapu/chra Say 11(1) 8 3 3 2 3 2 Appendix 2.2b. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3

S~ecies Ma~ Ma~ Ma~ Ma~Jun Jun Jun Jun Jul Jul Jul Jul Jul Au~ Au~ Au~ Au~ Se~ Se~Se~ Se~ Oct Family Leiodidae Agathidiumsp.1 4 2 11 3 4 Agathidiumsp.2 2 3 8 2 5 2 3 6 3 4 Agathidiumsp.3 3 3 6 1 Agathidiumsp.4 (2) 4 (1 ) 3 3 Agathidiumsp.5 5 3 2 2 Agathidiumsp.6 5 Agathidiumsp.7

Agathidiumsp.8 Agathidiumsp.9 2 Agathidiumsp.10 (1) Agathidiumsp.11 (1) Agathidiumsp.12 0\ Agathidiumdepressum FaU - (1 ) Agathidiumpulchrum LeConte Agathidium?oniscoides (2) Ag/yptinus/aevis (LeConte) Anisotornabasa/is (LeConte) (3) 2(3) 2(8) 2(7) 3(5) 2(1) 4(2) 3(4) 2(2) (2) (1 ) Anisotomab/anchardi (Hom) 1 2 7 9 6 16 1 1 Anisotomadisc%r (Melsheimer) 27(2) 32 31(4) 8(6) 12(13) 15(11) 20(6) 15(10) 31(4) 29(7) 5(1) 3 2 Anisotomageminata (Hom) Anisotomaglobososa Hatch (1) Anisotomahomi Wheeler 2 15(1) 40(2) 63(4) 60 68(1) 74 79 119 80(1) 31 25 4 4 6 3 5 Anisotornainops W.J.Brown 1(5) 2(1) (1) 3(1) 1(1) 5(4) (9) 8(6) 9(1) 1(2) (2) (3) (5) (3) Anogduspuritanus (FaU) 2 5 3 10 5 Anogdusobsoletus (Melsheimer) 3 3 3 2 2 Colenisimpunctata LeConte 2 2 3 3 2 2 Cyr/usasubtestacea (GyUenhal) Appendix 2.2b. continued

Isoplas/usfossor Hom Leiodescampbelli (Baranowski) Leiodespygmaea (Baranowski) Leiodessorenssoni (Baranowski)

Leiodessubülicomis (Baranowski) 2 2 5 2 Uocyrtusanigrie/avis (Hlisnikovsky)

UonothusfOfÜcomis Daffner 4 Triarthron pennsylvanicum Hom Family Mycetophagidae Myce/ophagusflexuosus Say 2(3) 2 2 3(2) (4) 1(1 ) 2(1) 3 4 2 2 2 01 N Myceotophaguspluripunctatus LeConte Mycetophaguspunctatus Say (2) (1 ) (1 ) 1 Myce/ophagusserrula/us Casey

Family Scaphidiidae ScaphidiumqUadriguttatum Melsheimer 3 2 1 Family Sphindidae Odon/osphindusdenücollis leConte 2 2 (1 ) Family Tenebrionidae BolitophaguscOfÜcola liliger Bolitophaguscomu/us (Panzer) Diaperismaculata Olivier

Neomidabicomis (Fabricius) 1 Platydemasp.A 1 (1) Upisceramboides Fabricius Appendix 2.2b. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Trogossitidae Tenebroides?cotticalis (Melsheimer) Thymalusmarginicollis Chevrolat

Family Zopheridae Phellopsisobcordata (Kirby)

0\ w Appendix 2.2e. Fungivorous Coleoptera eolleeted in flight-intereept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #3, Morrisburg, ON, (OG-3).

Eupariusmarmoreus (Olivier) 2 Euxenuspunctatus LeConte

Tropideresdorsalis (Thunberg)

Tropiderestricarinatus (Pierce) 1 2 4 Family Cerylonidae Geryloncastaneum Say

Gerylonunicolor (Ziegler) 3(1) 4 Philothermusglabriculus LeConte 0\ ..j::. Family Endomychidae Danaetestacea (Ziegler) Endomychusbiguttatus Say Lycoperdinaferruginea LeConte

Mycetinaperpulcra (Newman) 3 2 2 2 Phymaphorapulchella Newman 2 (1 ) Ranideaunicolor (Ziegler)

Family Erotylidae Dacnequadrimaculata (Say) (1) Triplaxdissimulator (Crotch) Triplax flavico/lis Lacordaire Triplax frosti Casey

Triplaxmacra LeConte 2 8 6(1) 2 2 Triplax thoracica Say Tritomamimetica (Say) 3 4 3 2 Tritomapulchra Say 2 6(1) 7 7 10 2 5 6 9(1) 4 2 2 Appendix 2.2c. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S~ecies Ma~ Ma~ Ma~ Ma~Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Se~ Se~ Se~ Oct Se~ Family Leiodidae Agathidiumsp.1 1 4 3 2 3 3 1 Agathidiumsp.2 1 1 2 2 4 4 Agathidiumsp.3 2 1 2 2 1 5 1 1 Agathidiumsp.4 2 1(1) 6 1 (1 ) 1 Agathidiumsp.5 2 8 2 1 Agathidiumsp.6 2 Agathidiumsp.7 1 Agathidiumsp.8

Agathidiumsp.9 1 2 3 2 Agathidiumsp.10

Agathidiumsp.11 (2) Agathidiumsp.12

0'1 Agathidiumdepressum Fan VI 1 1 (1 ) Aga/hidiumpu/chrum LeConte 4 2 2 3 3 2 3 3 Aga/hidium?oniscoides 2 Ag/yptinus/aevis (LeConte) Anisotornabasa/is (LeConte) 3(1) 4(1) 3 5(1) 5(1) 3(2) 8(2) 11 6 4 1(1) 2(1) 1 Anis%mab/anchardi (Hom) 5 3 7 15 9 2 4 8 2 2 Anis%madisc%r (Melsheimer) 26 5 14 26 17(1) 24 9(4) 29(3) 18(1) 13(1) 7 4 3 5 6 5 Anisotomageminata (Hom) Anis%maglobososa Hatch Anis%rnahomi Wheeler 1 7 26 79 103 71 53 141 158 135 160 100 69 35 32 16 7 8 3 2 Anisotornainops W.J.Brown 3(1) 2 1 2(3) 2(7) 4(3) 1(1) 1(1 ) 2(8) 2(1) (2) (1 ) Anogduspuri/anus (Fall) Anogdusobsoletus (Melsheimer) 4 3 1 3 Colenisimpunctata LeConte 2 5 3 2 2 6 4 9 7 4 2 8 8 3 Cyttusasub/es/acea (Gynenhal) 1 Appendix 2.2c. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Leiodidae (continued) Hydnobiusn. sp.

Hydnobiuslongidens LeConte 2 Isoplastusfossor Hom Leiodescampbelli (Baranowski) Leiodespygmaea (Baranowski) 2 3 435 3 4 34 11 4 Leiodessorenssoni (Baranowski) Leiodessubtilicomis (Baranowski) 20 53 54 26 13 14 17 23 25 10 14 22 11 20 43 11 5 Uocyrlusanigriclavis (Hlisnikovsky) 1 2 2 8 5 5 5 Uonothusforticomis Daffner

Triarthronpennsylvanicum Hom Family Mycetophagidae Mycetophagusflexuosus Say 2 13(1) 9 1 2 3(1) 3 7 3 4 ~ Myceotophaguspluripunctatus LeConte

Mycetophaguspunctatus Say 3 2 4 Mycetophagusserrulatus Casey

Family Scaphidiidae Scaphidiumquadriguttatum Melsheimer 2 2 Family Sphindidae Odontosphindusdenticollis leConte 2 2 Family Tenebrionidae Bolitophaguscorticola IlIiger Bolitophaguscomutus (Panzer) Diaperismaculata Olivier (1) Neomidabicomis (Fabricius) (1 ) Platydemasp.A

Upisceramboides Fabricius Appendix 2.2c. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Trogossitidae Tenebroides ?corticalis (Melsheimer) (1 ) Thymalusmarginicollis Chevrolat

Family Zopheridae Phellopsisobcordala (Kirby)

0\ -..J Appendix 2.2d. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #1, Lancaster, ON, (MF-l).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Anthribidae Choragussayi LeConte Eupariusmarmoreus (Olivier) Euxenuspunctatus LeConle Tropideresdorsa/is (Thunberg)

Tropiderestricarinatus (Pierce) 3 2 3 Family Cerylonidae Cetyloncastaneum Say 5 Cery/onunic%r (Ziegler) (1) 5(1) 2 2 Phi/othermusg/abricu/us LeConle 01 4 00 Family Endomychidae Danaetestacea (Ziegler) Endomychusbiguttatus Say (1 ) 1 Lycoperdinafenuginea LeConte 4 Mycetinapef/Julcra (Newman) 15(1) 23 5 7 3 10 2 4 Phymaphorapulchel/a Newman Ranideaunic%r (Ziegler)

Family Erotylidae Dacnequadrimacu/ata (Say) Trip/axdissimu/ator (Crotch) Trip/ax flavicol/is Lacordaire (2) Trip/ax frasti Casey Trip/axmacra LeConte Trip/ax thoracica Say (14) Tritomamimetica (Say) 2 Tritomapu/chra Say 2 2(1) 1(1) 2 4 3 Appendix 2.2d. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Seecies Ma~ Ma~ Ma~ Jun Ma~ Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug See See See See Oct Family Leiodidae Agathidiumsp.1 1 1 2 2 8 12 16 7 19 2 5 Agathidiumsp.2 5 2 10 15 3 4 8 1 11 1 2 Agathidiumsp.3 3 6 1 2 2 Agathidiumsp.4 (1) 3(1) (1) (1) 1(1) (2) Agathidiumsp.5 2(1) 6 1 2 Agathidiumsp.6 1 Agathidiumsp.7 2 (1) Agathidiumsp.8 3 Agathidiumsp.9 3 1 7· 5 5 6 Agathidiumsp.10

Agathidiumsp.11

Agathidiumsp.12 1 01 Agathidiumdepressum FaU \0 1 2 Agathidiumpu/chrum LeConte 2 Agathidium?onisCOides (1) Ag/yptinus/aevis (LeConte) Anisotomabasa/is (LeConte) 2 (15) 4(2) 4(4) 2(10) 2(1) 3(3) 6(2) 6(5) 2(6) 3 2 Anisotornab/anchardi (Hom) 2 3 14 7 3 7 19 19 Anisotomadisc%r (Melsheimer) 2 15(2)23(1) 19(11) 9(3) 19(6) 12(1) 10 25(2) 5 2(1) 5 2 2 Anisotomageminata (Hom) 4 4 Anisotomaglobososa Hatch 1 Anisotomahomi Wheeler 4 21 42(7) 60(4)86(1) 33(1) 64(4) 89(4) 72(1) 36(1) 31(6)16(3) 6(1) 3 7 3 Anisotomainops W.J.Brown 1(1) (4) 1 (4) (1) (1) (1) (8) Anogduspuritanus (Fan) 4 6 3 8 1 4 AnogdusobsoJetus (Melsheimer) 3 11 13 13 6 10 2 CoJenisimpunctata LeConte 2 2 5 2 6 7 3 3 3 2 Cyrlusasubtestacea (Gyllenhal) Appendix 2.2d. continued

Leiodeseampbelli (Baranowski) 2 Leiodespygmaea (Baranowski) 2 2 Leiodessorenssoni (Baranowski)

Leiodessubtilicomis (Baranowski) 3 3 9 5 3 5 3 5 2 2 1 Lioeyrlusanigrielavis (Hlisnikovsky) 1 3 2 Lionothusfortieomis Daffner 4 Triarthronpennsylvanieum Hom Family Mycetophagidae Myeetophagusffexuosus Say 3 3(1) 2(2) 1(1) 6(1) 5(1) 2(2) 8(1) 3 4 3 14 10 6 7 o...... :J Myeeotophaguspluripunctatus LeConte Mycetophaguspunctatus Say 2 2 Myeetophagusserrulatus Casey

Family Scaphidiidae SeaphidiumqUadriguttatum Melsheimer 1 4 5 Family Sphindidae Odontosphindusdentieollis LeConte 2 .ill Family Tenebrionidae BolitophaguscorticoJa liliger (1 ) Bolitophaguscomutus (Panzer) (1 ) (1) (1) Diaperismaculata Olivier (1 ) Neomidabicomis (Fabricius) PlatydemaspA

Upiseeramboides Fabricius Appendix 2.2d. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Trogossitidae Tenebroides?corlicalis (Melsheimer) Thymalusmarginicollis Chevrolal

Family Zopheridae Phellopsisobcordala (Kirby)

-...l ...... Appendix 2.2e. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #2, Bainsville, ON, (MF-2).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Anthribidae Choragussayi LeConte (1) Eupariusmannoreus (Olivier) Euxenuspunctatus LeConte Tropideresdorsalis (Thunberg) Tropiderestricarinalus (Pierce)

Family Cerylonidae Geryloncastaneum Say 1 (1) 2 Gerylonunicolor (Ziegler) 1(1 ) 5 Philothermusglabriculus LeConte -....J N Farnily Endomychidae Danaetestacea (Ziegler) 1 Endomychusbiguttatus Say 1(1) 2 (2) 2 Lycoperdinaferruginea LeConte 2 2 2 1 Mycetinaperpulcra (Newman) 7 2 4 6 4 2 1 Phymaphorapulchella Newman (1 ) Ranideaunicolor(Ziegler)

Family Erotytidae Dacnequadrimaculata (Say) Triplaxdissimulator (Crotch) Triplax flavicollis Lacordaire Triplax frosti Casey

Triplaxmacra LeConte Triplax thoracica Say Tritomamimetica (Say)

Tritomapulchra Say 3 6 2 4 2 2 Appendix 2.2e. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 SQecies Ma~ Ma~ Ma~Ma~ Jun Jun Jun Jun Jul Jul Jul Jul Jul AU9 AU9 AU9 AU9 SeQ SeQ SeQ SeQ Oct Family Leiodidae Agathidiurnsp.1 4 1 10 20 27 9 20 3 2 2 10 Agathidiurnsp.2 3 15 2 9 8 15 5 1 12 3 5 Agathidiurnsp.3 1 3 3 2 3 Agathidiurnsp.4 1(1) 2 Agathidiurnsp.5 5 5 5 4 Agathidiurnsp.6 1 Agathidiurnsp.7 2 2 3 2 Agathidiurnsp.8 1 Agathidiurnsp.9 2 3 2 2 3 Agathidiurnsp.10

Agathidiurnsp.11 (1 ) (1 ) Agathidiurnsp.12 2 3 4 -....J w Agathidiurndepressurn FaU Agathidiurnpulchrurn LeConte 2 Agathidiurn?oniscoides Ag/yptinuslaevis (LeConte) Anisotornabasalis (LeConte) 1(11 ) 2(2) 5(6) 4(7) 5(1) 10(4) 15(6) 14(6) 15(10) 7(1) 3 (1) Anisotornablanchardi (Hom) 9 1 1 21 17 46 39 1 Anisotornadiscolor (Melsheimer) 1(1) 16(1) 3(8) 4(4) 4 4 2 4 2 3 2 3 Anisotornagerninata (Hom) 2 Anisotornaglobososa Hatch 1 Anisotornahorni Wheeler 5 27 43(4) 85(3) 83(1) 89(1) 65 123(1) 92 78 48 27 9 6 6 11 1 Anisotornainops W.J.Brown (1 ) 1 1(4) Anogduspuritanus (FaU) 2 5 8 15 46 6 8 1 Anogdusobsoletus (Melsheimer) 1 3 1 2 3 1 6 1 2 1 Colenisirnpunctata LeConte 8 2 2 2 6 5 7 3 14 19 8 16 8 5 2 6 1 Cyrlusasubtestacea (GyUenhal) Appendix 2.2e. continued

Leiodespygmaea (Baranowski) 2 1 Leiodessorenssoni (Baranowski) 3 2 3 Leiodessubti!icomis (Baranowski) 3 2 3 2 2 4 7 3 2 3 2 2 Uocyrlusanigriclavis (Hlisnikovsky) Lionothusforlicomis Daffner

Triarlhronpennsy/vanicum Hom Family Mycetophagidae Mycetophagusflexuosus Say (2) 1(1) (1) 2(10) (1) ~) 3 5 4 2 8 5 20 7 7 4 -...l Myceotophaguspluripunctatus LeConte ~ (1 ) Mycetophaguspunctatus Say (1 ) 1(1) 2 3 2 Mycetophagusserrulatus Casey

Family Scaphidiidae Scaphidiumquadriguttatum Melsheimer 2 4 Family Sphindidae Odontosphindusdenticollis LeConte (1 ) Family Tenebrionidae Bolitophaguscorticola IlUger Bolitophaguscomutus (Panzer) Diaperismaculata Olivier Neomidabicomis (Fabricius) Platydemasp.A

Upisceramboides Fabricius Appendix 2.2e. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Trogossitidae Tenebroides?corocalis (Melsheimer) (1) Thymalusmarginicollis Chevrolat (1) Family Zopheridae Phellopsisobcordata (Kirby)

-....) V\ Appendix 2.2f. Fungivorous Coleoptera collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #3, Bouck's Hil1/Williamsburg, ON, (MF-3).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Seecies Ma~ Ma~ Ma~ Ma~Jun Jun Jun Jun Jul Jul Jul Jul Jul AU9 AU9 AU9 AU9 Sel:! Sel:! Sel:! Sel:! Oct Family Anthribidae Choragussay; LeConle Eupariusmarmoreus (Olivier) (1) (1 ) Euxenuspunctatus LeConte Tropideresdorsa/is (Thunberg) Tropiderestricarinatus (Pierce)

Family Cerylonidae Cery/oncastaneum Say (1 ) Cery/onunic%r (Ziegler) (1) (1 ) 1(2) Philothermusg/abricu/us LeConle 0\-...J Family Endomychidae Danaetestacea (Ziegler) Endomychusbiguttatus Say 1(1) 2 3 2 2 Lycoperdinaferruginea LeConte Mycetinaperpu/cra (Newman) (1 ) 5 3 6 11 7 7 Phymaphorapu/che/la Newman 2 Ranideaunic%r (Ziegler)

Family Erotylidae Dacnequadrimacu/ata (Say) (1 ) Trip/axdissimu/ator (Crotch) Trip/ax flavico//is Lacordaire Trip/ax frosti Casey Trip/axmacra LeConte 2 2 Trip/ax thoracica Say 2 1 2 Tritomamimetica (Say) 5 2 3 3 5 4 Tritomapulchra Say 2(1) 4 5 2 2 4 8 8 3 Appendix 2.2f. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S!;!ecies Ma~Ma~ Ma~ Ma~ Jun Jun Jun Jun Jul Jul Jul Jul Jul AU9 AU9 AU9 AU9 Se!;! Sep Se!;! Se!;! Oct Family Leiodidae Agathidiumsp.1 9 8 7 10 34 24 17 2 Agathidiumsp.2 2 3 10 4 4 9 7 Agathidiumsp.3 Agathidiumsp.4 1(1) 2 4 2 Agathidiumsp.5 3 6 2 2 Agathidiumsp.6 Agathidiumsp.7 Agathidiumsp.8 Agathidiumsp.9 6 Agathidiumsp.10

Agathidiumsp.11 (1 ) (1) Agathidiumsp.12 -..,J -..,J Agathidiumdepressum Fall Agathidiumpu/chrum LeConte 5 3 Agathidium?oniscoides (1 ) Ag/yptinus/aevis (LeConte) Anisotomabasa/is (LeConte) 4 2(5) 4 7 5 4 7 4 4 5(1) (1) 2 Anisotornab/anchardi (Hom) 7 4 2 1 10 5 24 7 Anisotornadisc%r (Melsheimer) (1 ) 58(1) 9(2) 3(7) 5(3) 11(1) 18(1) 4 5 Anisotomageminata (Hom) Anisotomag/obososa Hatch (4) (1 ) 1 Anisotomahomi Wheeler 5(1) 8(1) 50(2) 58(2) 15(1) 52 64 70 135 114(1) 106 90(1) 40 10 5 2 3 Anisotomainops W.J.Brown 9(15) 3(3) 1(2) 1(2) 5(5) 4 5(6) 2(6) 2(5) 1(3) 2(9) 3(3) 1(2) 2(2) (1 ) Anogduspuritanus (Fall) 2 3 15 4 1 Anogdusobsoletus (Melsheimer) 1 6 10 5 5 1 2 Colenisimpunctata LeConte 2 4 12 13 9 16 7 14 15 10 8 2 3 2 Cyrlusasubtestacea (Gyllenhal) Appendix 2.2f. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Family Leiodidae (continued) Hydnobiusn. sp. 1 1 3 2 2 Hydnobiuslongidens LeConte 2 5 2 2 3 5 3 2 Isoplastustossor Hom 6 21 7 1 2 5 Leiodescampbelli (Baranowski)

Leiodespygmaea (Baranowski) 2 2 2 2 Leiodessorenssoni (Baranowski) 2 1 Leiodessubülicomis (Baranowski) 3 5 5 8 3 2 1 4 5 2 3 5 2 UocyrtusanigricJavis (Hlisnikovsky) 2 2 3 6 316 16 6 6 12 2 Uonothusforticomis Daffner 4 11 2 13 8 2 5 3 3 Triarthronpennsylvanicum Hom Family Mycetophagidae Mycetophagusflexuosus Say 3 (1 ) 1(1) 5 2 (1 ) 2 2(1) 9 4 3 -.....l 00 Myceotophaguspluripunctatus LeConte Mycetophaguspunctatus Say 2 2 1(1) (2) 3 MycetophagussefTUlatus Casey

Family Scaphidiidae Scaphidiumquadriguttatum Melsheimer 2 1 2 2 2 1 m 5 Family Sphindidae Odontosphindusdenticollis LeConte (5) Family Tenebrionidae Bolitophaguscorticola lIIiger Bolitophaguscomutus (Panzer) Diaperismaculata Olivier

Neomidabicomis (Fabricius) PlatydemaspA (1) Upisceramboides Fabricius Appendix 2.2f. continued

-.....) \0 CONNECTING STATEMENT The comparative biodiversity study discussed in Chapter 2 examined eleven families ofprimarily fungivorous Coleoptera. One of the families not considered in that study was the Nitidulidae. Nitidulids, or sap beetles, specialize on a wide v ari et y ofhosts (i.e., plants, carrion, fungi, sap) for feeding and breeding. Because they are not strictly fungivorous they wère not inc1uded in the analysis of the fungivorous Coleoptera and a separate analysis was performed. While much information has been compiled on the biology of a few nitidulid species considered minor agricultural pests (e.g., Glischrochilus quadrisignatus), little is known about the ecology and distribution of the majority of other sap beetles in eastem Ontario. This study was one of the first to provide an unbiased comparative biodiversity survey ofthe nitidulid species present in small forest stands in North America.

80 CHAPTER 3 - Sap beetles (Coleoptera: Nitidulidae) in managed and 'old-growth' forests in south-eastern Ontario, Canada

ABSTRACT Nitidulid beetles were sampled from 'old-growth' and mature-managed hemlock hardwood forests in southeastem Ontario. Large-area flight-intercept traps (FITS) and trunk-window traps (TTs) were operated for 22 weeks in 2003, and yielded a total of 2,129 nitidulid beetles comprising 30 species. Species abundance was higher in managed forests and species richness was higher in 'old-growth' forests. These results, however, were strongly influenced by the dominance of Glischrochilus quadrisignatus (Say) at two of the six forest sites. Pal/odes pallidus (Palisot de Beauvois) was stronglyassociated with managed forests. Glischrochilus sanguinolentus (Olivier) was coUected much more frequently in TTs than in FITs and data suggested a possible association ofthis beetle with 'old-growth' forests. Pal/odes pallidus and Cychramus adustus Erichson, both known to feed on fleshy white fungi, displayed a c1ear division in seasonal abundance peaks, suggesting that resource partitioning may be occurring.

81 INTRODUCTION Anthropogenic disturbances such as logging, agriculture, fire suppression, and urban development are responsible for the loss and fragmentation of forests globally. In the temperate mixed-wood forests of eastem Ontario (the Great Lakes-St. Lawrence Region or GLSL), disturbances stemming from human settlement between the mid-1700s and early 1900s destroyed much of the original forest (Suffling et al. 2003). Forests in the southem part of the GLSL are highly fragmented with little connectivity between forest stands and little core habitat in remaining stands. The region is heavily populated and natural areas have been greatly modified by agriculture and urbanization, making conservation of the area's remaining old-growth forests an urgent priority. Comparative biodiversity studies have received much attention in recent years and a large volume ofwork has been published on the diversity ofvarious taxa in managed and unmanaged (old-growth) forests across the globe. Coleoptera especially have received a lot of attention in forest diversity and management studies because they are thought to be particularly useful for biodiversity and environmental assessment work for several reasons: the taxonomy and distribution of many families are well-known in sorne geographic regions; they are easy to preserve, store and prepare for later identification (Hammond 1997); and, they are one of the most ecologically diverse insect orders (Grove 2002). Old-growth forests host several rare and/or specialist beetle species (Siitonen and Martikainen 1994; Berg et al. 1994) and the effects offorest management on saproxylic beetle fauna have been widely studied (e.g., Chandler and Peck 1992; Martikainen et al. 2000). Most of these studies have linked the diversity and abundance of saproxylic beetles to several variables, inc1uding the volume of dead wood present in the forest (Schiegg 2000; Simila et al. 2003), the diversity (species, size and decay class) of dead wood (0kland et al. 1996), the abundance and diversity ofwood-inhabiting fungi (Kaila et al. 1994; Komonen 2003), and the degree offorest disturbance (Vaisanen et al. 1993; Siitonen and Martikainen 1994). Sorne species (e.g., bark beetles) can benefit from forest management and freshly dead wood (Hammond et al. 2001); however, others are thought to be severely affected by forest disturbance.

82 The majority of forest arthropod research has taken place in northern Europe and relatively few studies have been conducted in North America (but see Chandler 1991, Chandler and Peck 1992, Werner and Raffa 2000, Buddle et al. 2000, Hammond et al. 2004). There is little known about the response of the Canadian arthropod fauna to forest management (Hammond 1997) and very little published information on the effects of forest management on beetle diversity is available for the mixed northern hardwood forests ofthe GLSL region (Werner and Raffa 2000). Zeran et al. (2004) studied the impact offorest management on fungivorous Coleoptera. One of the families not considered in that analysis was the Nitidulidae. Although some members of the Nitidulidae are fungivorous, many species within the family feed and breed on very different hosts (i.e., sap, flowers, carrion, decayed plant matter) and thus merit a separate analysis. While the family as a whole is not considered to be saproxylic, many species are dependent on forest habitat during at least some part of their life cycle. Sap beetles (Nitidulidae) are small beetles usually ranging in size from 1.5 to 12 mm (Habeck 2002). Although they are primarily saprophagous and mycetophagous, the habits of nitidulids are quite variable, with species being found on flowers, fruits, sap, fungi, decaying/fermenting plant tissues or carrion (Habeck 2002). Some species are considered to be minor agricultural pests, including Carpophilus lugubris Murray on corn, Glischrochilus quadrisignatus (Say) on corn, and Stelidota geminata (Say) on strawberries and raspberries (Dowd and Nelsen 1994; Blackmer and Phelan 1995). Glischrochilus quadrisignatus is also considered to be a primary vector ofthe oak wilt pathogen, Ceratocytisfagacearum (Bretz) (Cease and Juzwik 2001). Nitidulid beetle make up a small but interesting component of forest arthropod populations, however, few studies have looked at them outside of agriculturallandscapes. The objectives ofthis study were to: (1) conduct a faunal inventory and document the diversity, distribution and phenology of nitidulid beetles in selected forest ecosystems of southeastern Ontario; and (2) compare the diversity ofnitidulids between managed and 'old-growth' forests. Because old-growth forests usually support a more diverse arrayof microhabitats than do managed forests, it was predicted that nitidulid beetles would be more diverse in 'old-growth' forests.

83 MATERIALS AND METHODS Study Sites Six forest stands were selected in southeastem Ontario (Table 3.1, Figure 3.1), in section L.2 of the Great Lakes-St. Lawrence Forest Region (Rowe 1972). Three forests were managed (MF-l, MF-2, MF-3) and three were considered 'old-growth' (OG-1, OG-2, OG-3). For this study 'old-growth' forests were considered those which are ofsufficient age (averaging over 120 years of age), demonstrate many 'old-growth characteristics' (e.g., pit and mound topography, large trees, large volumes of coarse woody debris, multi-aged stand structure), and have remained mostly free ofhuman disturbance for at least several decades (e.g., Stabb 1996). Managed forests also demonstrated many 'old grwoth' characteristics, only having slightly lower volumes of coarse woody debris and a less distinct pit and mound relief. AlI six forest stands were relatively small (between 5.9 ha and 13.8 ha), fragmented remnant forests, with similar elevation, topography, soil, and c1imate characteristics. AlI sites were dominated by eastem hemlock (Tsuga canadensis (L.) Carr.), with lesser quantities of sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifolia Ehrh.), white pine (Pin us strobus L.) and a variety of other hardwoods. Managed forests ranged in age from 100-150 years of age, while 'old growth' forests were aIl over 120 years of age (Table 3.1). In the fall of1999, about one third of the total wood volume was removed from all three of the managed sites by the local forest company (P. Wensink, Domtar Inc., 2003, personal communication).

Sample Collection Coleoptera were collected using two trap types. A large-area flight-intercept trap (FIT) was based on the design ofPeck and Davies (1980). Two 1.85 m pieces ofrebar were inserted into the forest floor approximately 2 m apart. A 1.25 m x 1.85 m mesh panel of standard black window screening was stretched taut between the upright pieces of rebar. Traps were stabilized by tying nylon cord from the top of each piece of rebar to nearby trees. A wooden trough (1.85 m x 0.5 m x 9 cm) lined with thick c1ear plastic was placed under the mesh panel. A c1ear plastic rain roof (2.36 m x 1.5 m) was stretched over the trap and tied to nearby trees.

84 A modified version of the trunk-window trap (TT) described by Kaila (1993) was also us.ed. A transparent plexiglass window (23 cm x 14 cm) was attached above an inverted 2L pop bottle with the bottom cut away. The traps were fastened to trees with nylon cord at about 1.3 m above ground. Because only a small portion of the funnel was in direct contact with the tree trunk the traps targeted flying insects more than those crawling on the trunk. Trunk traps were placed randomly in each plot within 25 m of the FIT, on dead trees with polypore fungus. Iftrees with fungus could not be located, traps were placed on dead trees (or, in one case, on moribund trees, since dead trees could not be located). Traps were placed on multiple tree species in a variety of decay classes, since the goal of the study was to assess the nitidulid diversity ofthe entire stand and not just of a particular tree species. Two plots approximately 30 m radius were established in each forest site, with one FIT and three TTs in each plot (12 FITs and 36 TTs in total). A saturated salt water solution was used for the collecting fluid in all traps. Traps were operated from 29 April 2003 until3 October 2003 (22 weeks). AlI traps were emptied weekly throughout the season using a modified aquarium net (FITs) or a kitchen strainer (TTs) and collecting fluid was replaced as it evaporated.

Nitidulid Identification Beetles were initially stored in 70% ethanol and later mounted on points and identified to species. Nitidulid specimens were identified to genus using standard references (Downie and Arnett 1996; Arnett et al. 2002). Although the subfamily Cateretinae is now considered to be a separate family (Brachypteridae) (Habeck 2002), species from this group (e.g., Brachypterus spp. and Kateretes spp.) were included in the Nitidulidae for this study. Species identifications were made using keys and descriptions in taxonomic revisions or by collaborating specialists (A. Cline, Louisiana State Arthropod Museum). Species identifications were verified by consultation with specialists or by comparison to identified specimens in the Canadian Museum of Nature, Aylmer, QC (CMN) and the Lyman Entomological Museum, McGill University, Ste Anne-de-Bellevue, QC (LEM). Voucher specimens are deposited in LEM and CMN.

85 Data Analyses Species abundance data from each trap were pooled throughout the season, resulting in 48 samples (12 FIT samples, 36 TT samples). The number of species observed (raw species richness) in each site was recorded. Rarefaction was used to estimate the expected species richness in each site. Rarefaction can also be considered a diversity index because it accounts for both species richness and abundance (Hammond et al. 2004; Olszewski 2004). Standard deviations were plotted for each point on the rarefaction curve. For rarefaction calculations, data from FITs and TTs were pooled by site, resulting in six "samples". Rarefaction was performed using ECOSIM 7.0 (Gotelli and Entsminger 2001). Fixed, individual-based sub-samples (at increments of20) were used in the calculation and the data matrix was randomized 1000 times. To examine

evenness and dominance of the data, Fisher's ct. and Simpson's indices were calculated

using EstimateS 6.0bl (Colwe1l2001). Fisher's ct. is one ofthe most robust diversity measures and is relatively unaffected by sample size (Magurran 2004). Because of the small sample size, Simpson's index was chosen as the most appropriate measure of dominance. For these calculations, FITs and TTs were treated as individual samples (2 FITs and 6 TTs for each site). {j-diversity was calculated for each pair of sites (with

pooled data from both FITs and TTs) using the Morisita-Hom index (CMH). This index was chosen as a similarity measure because it is strongly influenced by species richness and sample size (Wolda 1981; Magurran 2004). Because the data did not display enough variation (i.e., a low number oftraps collected a large number ofspecies and specimens) an ordination could not be performed. Instead, cluster analysis was performed on the data from FITs to compare the similarity ofbeetle assemblages between the twelve traps. A Sorenson distance measure and a group averaging linkage method were used in PC-Ord 4.0 (McCune and Mefford 1999) to mn the cluster analysis. Indicator species analysis (Dufrêne and Legendre 1997) was also conducted using the pro gram PC-Ord 4.0 to determine ifbeetle species showed significant associations with forest type. Indicator species analysis uses relative abundance and proportional frequency of each species in each sample and produces an indicator value (lndVal) for each species ranging from 0 (no association) to 100 (extremely strong association). An IndVal of 100 means that the presence of a species

86 points to a particular habitat (or group) without error (McCune et al. 2002; Hammond et al. 2004). Rare species (with only a few occurrences) do not yield an IndVal higher than expected by chance (McCune et al. 2002). Indicator values were tested for significance using a Monte Carlo randomization (with 1000 permutations) which resulted in a p-value for each IndVal (McCune et al. 2002; Hammond et al. 2004). Nitidulid catches in TTs were infrequent and patchy, thus, c1uster analysis and indicator values were calculated only for FIT data.

RESULTS Richness, Abundance and Phenology A total of2,129 nitidulid beetles comprising 30 species was collected. Complete species lists for each forest site with the number of specimens in each trap type in each week are in Appendices 3.1a to 3.1f. Traps in managed forests collected over twice as many individuals (1501) than those in 'old-growth' forests (628). This difference was mainly due to the number of specimens of Glischrochilus quadrisignatus collected in managed forests (51 % of all nitidulids collected in managed forests were G. quadrisignatus). The total number of species did not differ greatly between forest types, with 24 species occurring in 'old-growth' and 25 species in managed forests. The relative abundance of Glischrochilus quadrisignatus was highest in May, in late July and in mid to late September (Fig. 3.2a). Glischrochilus sanguinolentus numbers were highest in May and steadily dec1ined untillate July, after which the species was rarely collected (Fig. 3.2b). Cychramus adustus Erichson peaked in early July while numbers of Pallodes pallidus (Palisot de Beauvois) were highest from early August until late September (Fig. 3.3a). Omosita colon (L.) was found at the beginning of the season (May to July) but was rarely collected after mid-July (Fig. 3.3b). The five most commonly collected species of Epuraea (E. ru/a, E. labilis, E. erichsoni, E. depressa, and E. rufida) were caught from mid-May until mid-August, after which no more specimens were collected (Fig. 3.4).

87 Site Comparisons

The Fisher's CI. and Simpson's indices for each site generally ranked 'old-growth' sites as having higher diversity than managed sites (Table 3.2). However, one managed site (MF-3) had diversity values more similar to those of the three 'old-growth' forests, than to the other two managed forests. The Simpson's index indicated much lower diversity for two managed sites (MF-l and MF-2) than any other, reflecting the dominance of G. quadrisignatus at these two sites. The mean value of Fisher's CI. for 'old growth' was 4.31 and for managed forests, 3.79. The mean value of the Simpson's index was 5.20 for 'old-growth' and 3.33 for managed forest. However, site MF-3 may have skewed the results for managed forests when it was averaged together with the other two sites in this way. Rarefaction-estimated species richness for each site (compared at 120 individuals) is presented in Table 3.2 and rarefaction curves are in Fig. 3.5a. Standard deviations for each rarefaction estimate are also presented, allowing for an indication of the significance of each comparison. On a site by site basis, significant differences were as follows: OG- 1 was significantly higher than MF-l and MF-2; OG-2 was significantly higher than MF- 2; and OG-3 was significantly higher than all managed sites. A second rarefaction based on pooled managed versus 'old-growth' sites (compared at 620 individuals) showed that 'old-growth' sites differed significantly from managed forests (Fig. 3.5b). Pairwise Morisita-Hom values demonstrated several differences in {3-diversity. Sites MF-l and MF-2 had an almost identical species composition, with a similarity value of 0.990. MF-l and MF-2 were not similar to the third managed site (MF-3), with {3- diversity values of 0.396 and 0.352 respectively. The three 'old-growth' sites also varied in their similarity to each other (0.496 for OG-3 and OG-l to 0.814 for OG-3 and OG-2). There was high {3-diversity between the three 'old-growth' sites and the three managed sites (CMH values ranging from 0.226 to 0.446). {3-diversity for pooled managed versus 'old-growth' sites was 0.331.

Species Composition The c1uster analysis indicated a c1ear difference in the overall species composition ofnitidulids between 'old-growth' and managed forests with the managed sites and the

88 'old-growth' sites diverging very early in the dendrogram (Fig. 3.6). Results from the indicator species analysis of nitidulids collected with flight-intercept traps are presented in Table 3.3. Only two species, Glischrochilus quadrisignatus and Pal/odes pal/idus, showed a significant association with either forest type, both species being strongly associated (p ~.05) with managed forests (Table 3.3). The mean number ofindividuals of G. quadrisignatus was 256.33 ± 96.38 for managed stands and 6.67 ± 2.08 for 'old growth'. The mean number of P. pal/idus individuals collected was 24.67 ± 3.69 for managed and 1.00 ± 0.50 for 'old-growth'.

Effects of Glischrochilus quadrisignatus Because G. quadrisignatus caused large variation between managed sites, data sets were modified to exclude this species and analyses were re-mn on data from all sites to investigate the effect that this beetle had on 'old-growth'/managed forest comparisons.

When G. quadrisignatus was excluded, the Fisher's Cl and Simpson's indices ranked managed forests as having a slightly higher diversity than 'old-growth' forests. Rarefaction-estimated species richness indicated that there was no significant difference between 'old-growth' and managed forests; however, due to the lower sample sizes (1340 individuals), such conclusions may be prematur~. Cluster analysis indicated a similar trend, with sites and traps not distinctly separated from each other.

DISCUSSION Richness, Abundance and Phenology of Nitidulidae

Fisher's Cl, Berger- Parker and rarefaction deal with Cl-diversity, the diversity of species within a community or habitat (Southwood 1978). Rarefaction, while providing un-biased estimates of species richness at each site, also allows for standard deviations to be calculated for each estimate, facilitating a comparison between Cl-diversity in each forest site and between each habitat. In contrast, (j-diversity is a measure of the rate and extent of change in species along a gradient from one habitat to another (Southwood 1978). Distributions ofpairwise (j-diversity measures (such as those generated by the Morisita-Horn index) can be compared directly (Magurran 2004). Cluster analysis is another intuitive, simple method to compare (j-diversity by graphically representing

89 differences amongst samples/sites (Magurran 2004). Sites or samples that cluster together are more similar to one another than to those that are located at a distance from eachother. Sap beetles were much more abundant in managed forests (1501 individuals) than in 'old-growth' forests (628 individuals). Rarefaction-estimated species richness was higher in 'old-growth' sites than in managed sites and the Simpson's and Fisher's ex diversity indices indicated a similar pattern. Rarefaction curves (Fig. 3.5) indicated higher diversity in 'old-growth' than in managed sites; however, the lines for two sites are very short, indicating that sampling was probably not sufficient to produce a clear picture of the nitidulid species richness in those sites. Glischrochilus quadrisignatus represented 37% ofthe nitidulids collected from all six sites. The second most abundant species was Omosita colon, comprising almost 20% ofthe total nitidulid catch in aIl sites. In contrast, approximately 47% ofthe species collected (14 of30 species) were represented by less than ten individuals. One of the problems associated with trap data is that many species are wide-ranging 'tourists' that can sometimes be found in habitats where they do not reproduce or develop (NiemeUi. 1997). Often it is difficult to make such conclusions, especially iflittle is known of the biology of each species. In this study, no attempt was made to draw conclusions on individual habitat preference for rare species. A clear division in seasonal abundance of C. adustus and P. pallidus was observed (Fig. 3.3a), with C. adustus numbers peaking from mid-June to late July, and P. pallidus abundance being highest from late July through until the end ofSeptember. Because P. pallidus and C. adustus have similar feeding and breeding requirements (Crowson 1984; Lawrence 1998), this seasonal pattern probably indicates that resource partitioning is occurring. Both Omosita c%n and Omosita discoidea were abundant at the beginning of the season (May to July) but were rarely collected after mid-July. Both species of Omosita feed on carrion and this seasonal distribution may be related to the low abundance of carrion late in the season, or with avoidance of resource competition with other carrion feeding Coleoptera. Such resource partitioning among carrion-feeders has been previously suggested within the Leiodidae (Peck and Anderson 1985) and the Silphidae (Anderson 1982). There were three clear peaks of abundance in the seasonal

90 distribution of Glischrochilus quadrisignatus: in May, in late July, and again in mid September (Fig. 3.2a). This seasonal pattern was similar to that observed by Dowd and Nelson (1994) for G. quadrisignatus and suggests that this species has two generations per year. The species overwinters as adults (Luckman 1963), accounting for the high peaks in spring and faH. Several species were coHected only during relatively short periods oftime (e.g., Epuraea n. sp. only in June). Such patterns stress the importance of season-Iong sampling, since the species might be missed if sampling occurs only for a short time or at widely space intervals.

Comparison of Forest Types In accordance with our prediction, 'old-growth' forests did support higher species diversity than managed forests. There was, however, a confounding variable. One managed forest (MF-3) consistently had higher species diversity than the other two managed sites. These differences were mainly due to the abundance of Glischrochilus quadrisignatus. This species showed a very strong association with managed forests. However, over 94% of G. quadrisignatus individuals collected in managed forests were found in two sites (MF-l and MF-2), with the third managed site (MF-3) only accounting for 5% of the catch. The presence of large numbers of G. quadrisignatus at only two of the six sites can probably be explained by the attraction ofthis species to maize crops (Luckman 1963; Blackmer and Phelan 1995). The two sites in which G. quadrisignatus was most abundant were the only two 'forests' surrounded by active corn fields; these fields likely acted as a bait to the beetles, greatly increasing their abundance in adjacent forested areas. Like G. quadrisignatus, Kateretes scissus was also collected onlyat sites MF-l and MF-2, albeit in much lower numbers. Pal/odes pallidus showed a strong association with managed forests. Unlike, G. quadrisignatus, however, this species was found in relatively even numbers in aIl managed forests, while rarely being collected in 'old-growth' sites (Table 3.4). Pallodes pal/idus is fungivorous and is widely distributed throughout the eastern two-thirds of North America (A. Cline 2004, personal communication). The species has been found in association with fleshy Basiodiomycetes fungi such as Amanita DilI.ex Boehm., Boletus Tourn.ex Adans, Pluteus Fr., and Russula Pers. (Leschen 1988) and its larvae are known

91 to feed and breed in basidiocarps (Crowson 1984; Lawrence 1998). The preference of P. pal/idus for fleshy mushrooms might explain its habitat preference (since these mushrooms tend to be more common on dead and decaying trees in hardwood forests), excePt that aIl six forest sites were dominated by hemlock (Tsuga canadensis), with the remaining tree species made up of a variety ofhardwoods (e.g., sugar maple and beech). In fact, 'old-growth' forests had more decaying hardwood than did managed forests. Thus, further investigation is necessary to determine the factors influencing the stronger association of P. pallidus with managed forests. Cychramus adustus was found almost exc1usively (95% of aIl specimens) in site MF-3 (Table 3.4). This may be due to geographic distribution. Site MF-3 was the most westerly (and northerly) located forest, foIlowed by OG-3 and OG-l (the only other two sites where the species was coIlected). The other three sites were located at the eastern end ofthe study range. Epuraea erichsoni Reitter was coIlected almost exc1usively (92% of aIl specimens) at site OG-l. Epuraea n.sp. was also found exc1usively (aIl 32 individuals) at site OG-l. Species such as this, which are associated with this particular 'old-growth' site, probably demonstrate a patchy distribution and may be relict populations resulting from the large-scale fragmentation that has occurred throughout the area. Site OG-l has been designated as a 'Demonstration Forest' by the Eastern Ontario Model Forest. Of aIl the sites, OG-l was perhaps the best representative of a healthy, mature/'old-growth' forest, and thus likely contains more micro-habitats and niches for species, particularly old-growth forest specialists. This site is potentially valuable for future biodiversity studies and would be ideal for investigation of fungus-insect relationships, especially for less commonly coIlected taxa such as Epuraea.

Resolving Patterns: Collecting Methods and Species Identification The most common method used to collect nitidulid beetles is baited (usually with fermenting fruits and vegetables or bread) traps (e.g., Keeney et al. 1994; Blackmer and Phelan 1995). In this study, baits were not used, and nitidulids were collected in traps (FITs, TTs) that are unbiased in that they do not deliberately attract certain species. This may have provided a more accurate picture of the nitidulid flight activity in the forests. While the majority ofnitidulids were collected with FITs in this study, 75% of the

92 Glischrochilus sanguinolentus specimens were collected from trunk-window traps. Of the four species of Glischrochilus collected, G. sanguinolentus is the species most commonly associated with fungi, especially bracket fungi such as Fomes fomentarius (L.ex Fr.) Kickx (Matthewman and Pielou 1971). Since trunk traps were placed on dead trees close to fungus, the dominance ofthis species in trunk traps may be explained by its attraction to basidiocarp fungi. However, because only beetles collected with flight intercept traps were used in the indicator species analysis, the association of G. sanguinolentus with 'old-growth' forests is under-estimated (e.g., bypooling FITs with trunk-traps, 73% of G. sanguinolentus individuals were collected in 'old-growth' forests, while only 27% were collected from managed sites). These results suggest that it would be beneficial if, when studying the Nitidulidae, a combination of trapping methods were used (i.e., flight-intercept traps, pitfall traps, bait stations, and fungus sampling). Studies that use only baited traps risk excluding species that specialize on flowers and fungus (e.g., G. sanguinolentus, E. depressa, and E. labilis). Several previous studies that conducted inventories or compared beetle assemblages have not identified individuals of the genus Epuraea to species (e.g., Kaila et al. 1994; Blackmer and Phelan 1995). The genus Epuraea has more North American species than any other genus ofNitidulidae, its members 'specializing' on a wide variety ofresources for both feeding and breeding. As previously mentioned, several species of Epuraea were found exclusively at certain sites. Indicator species analysis associated several species of Epuraea with either 'old-growth' or managed forests (Table 3.3); although the IndVals were not significant because of small sample sizes, c1earer associations might be seen with further collecting. Species-Ievel identification of genera like Epuraea in comparative biodiversity studies is necessary ifwe wish to understand how beetles (especially rare ones) respond to environmental variables. It is important to consider each species individually when trying to uncover ecological patterns since such patterns may be obscured if only the overall abundance or presence ofhigher taxa is considered (Sverdrup-Thygeson 2001).

93 Conclusion: Forest Management and Nitidulidae Initially, results indicated that forest management has a negative impact on the species richness of nitidulid beetles. However, when G. quadrisignatus was excluded, the difference was less apparent. This result is similar to that found by Blackmer and Phelan (1995) where the diversity of nitidulids in agricultural fields was found to be similar to that in adjacent woodlots. Zeran et al. (2004) found no difference in the diversity of fungivorous Coleoptera in managed versus 'old-growth' forests and discussed possible reasons for this result, one ofthe most intuitive being that the forest management occurring at these sites was not disruptive enough to cause large-scale differences in species richness between forests. Most studies that find such differences look at extreme forest management such as clearcuts (e.g., Buddle et al. 2000). Several species, however, were strongly associated with particular habitats (e.g., P. pal/idus, G. sanguinolentus, and various Epuraea species), indicating that while forest management may not have an effect on overall diversity of nitidulids, it can influence the species composition of forests. Further study is required to obtain more detailed data on the resource/host use by many species of nitidulids; only then can clear conclusions as to their habitat preference be made. Of course, as with any localized comparative study, the association ofbeetles in this study with a particular type of forest (either managed or 'old-growth') can, in reality, only be applied to forests in southeastern Ontario; similar studies in other North American regions (i.e., in the deciduous zone, the boreal zone, the coastal rainforests or even in the less disturbed areas of the GLSL forest region) would be required to determine how general these patterns are.

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96 Lawrence JF. 1998. Mycophagy in the Coleoptera: Feeding Strategies and Morphological Adaptations. pp 1-27 in N Wilding, NM Collins, PM Hammond, JF Webber (Eds), Insect-Fungus Interactions. Proceedings ofthe Ilh Symposium of the Royal Entomological Society ofLondon. London: Academic Press. Leschen RA. 1988. Pal/odes austrinus, a new species ofNitidulidae (Nitidulinae) with discussions on Pal/odes mycophagy. Journal ofthe New York Entomological Society 96: 452-458. Luckman WH. 1963. Observations on biology and control of Glischrochilus quadrisignatus. Journal ofEconomic Entomology 56: 681-686. Magurran A. 2004. Measuring Biological Diversity. Malden, USA: Blackwell Publishing. Martikainen P, Siitonen J, Punttila P, Kaila L, Rauh J. 2000. Species richness of Coleoptera in mature managed and old-growth boreal forests in southem Finland. Biological Conservation 94: 199-209. Matthewman, W.G. and Pielou, D.P. 1971. Arthropods inhabiting the sporophores of Fomesfomentarius (Polyporaceae) in Gatineau Park, Quebec. The Canadian Entomologist 103: 775-847. McCune B, Mefford M.J. 1999. PC-ORDo Multivariate analysis ofecological data, version 4. MjM Software Design, Gleneden Beach, Oregon. McCune B, Grace JB, Urban DL. 2002. Analysis of ecological communities. MjM Software Design, Gleneden Beach, Oregon. NiemeUi. J. 1997. Invertebrates and boreal forest management. Conservation Bi%gy Il: 601-610. 0kland B, Bakke A, Hagvar S, Kvamme T. 1996. What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southem Norway. Biodiversity and Conservation 5: 75-100. Olszewski TD. 2004. A unified mathematical framework for the measurement ofrichness and evenness within and among multiple communities. Oikos 104: 377-387 Peck SB, Anderson R.S. 1985. Seasonal activity and habitat associations ofadult small carrion beetles in southem Ontario (Coleoptera: Leiodidae: Cholevinae). The Co/eopterist 's Bul/etin 39: 347-353.

97 Peck SB, Davies AB. 1980. Collecting small beetles with large-area ''window'' traps. The Coleopterist's Bulletin 34: 237-239. Rowe JS. 1972. Forest Regions of Canada. Department ofFisheries and the Environment. Canadian Forestry Service, Publication No. 1300. Schiegg K. 2000. Are there saproxylic beetles species characteristic ofhigh dead wood connectivity? Ecography 23: 579-587. Siitonen J, Martikainen P. 1994. Occurrence ofrare and threatened insects living on decaying Populus tremula: A comparison between Finnish and Russian Karelia. Scandinavian Journal ofForest Research 9: 185-191. Simila M, Kouki J, Martikainen P. 2003. Saproxylic beetles in managed and seminatural Scots pine forests: quality of dead wood matters. Forest Ecology and Management 174: 365-381. Southwood T .R.E. 1978. Ecological Methods with particular reference to the study of insect populations. Second edition. Chapman and Hall, London. Stabb M. 1996. Ontario 's old growth: a learner's handbook. Ottawa, ON: Canadian Nature Federation. Suffling R, Evans E, Perara A. 2003. Presettlement forest in southern Ontario: ecosystems measured through a cultural prism. The Forestry Chronic/e 79: 485- 501. Sverdrup-Thygeson A. 2001. Can 'continuity indicator species' predict species richness or red-listed species of saproxylic beetles? Biodiversity and Conservation 10: 815-832). Vaisanen R, Bistrom 0, Heliovaara K. 1993. Sub-cortical Coleoptera in dead pines and spruces: is primeval species composition maintained in managed forests? Biodiversity and Conservation 2: 95-113. Werner SM, Raffa KF. 2000. Effects offorest management practices on the diversityof ground-occurring beetles in mixed northern hardwood forests of the Great Lakes Region. Forest Ecology and Management 139: 135-155. Wolda H. 1981. Similarity indices, sample size and diversity. Oecologia 50: 296-301.

98 Zeran RM, Anderson RS, Wheeler TA. 2004. Biodiversity of fungivorous beetles (Coleoptera) in managed and old-growth hemlock-hardwood forests in southeastem Ontario, in preparation.

99 Table 3.1. Location, tree species, size, age and general description ofstudy sites.

Site Location SEecies ComEosition* Size A~e General Information OG-l Cornwall,ON He37Mr3SBel7Pw90h2 9.3 ha 160-210 Private ownership. Stand likely 45°02.l60'N years selectively harvested about 160 years 74°47.470'W ago (high-grade - took only largest and best trees). Surrounded by sugar maple forest, city roads, and agricultural land. Acidic soil, fresh to moist; uElands sand~ loarn. OG-2 Lancaster, ON He3SBe3SMhlsPwloOhs -12 ha + 120 Owned by St. Lawrence Parks 45°07.750'N years Commission ("Raisin River Park", 74°30.875'W (climax operations closed 1992). Some forest) evidence of thinning in past. Surrounded mostly by Hwy 401 and the Raisin River. Flat, low-lying terrain. OG-3 Morrisburg, ON He30Be3oMh300h 10 - 8 ha + 130 Owned by St. Lawrence Parks 44°56.750'N years (late Commission ("Upper Canada Sugar 75°06.035'W succession Bush"/"Crysler Farm Battlefield Park al forest) Forest''). On west edge of 80 ha sugarbush; county roads and clearings to east, north and south of site. Mesic to wet; hummocky sandy loam over cla~Elain. MF-l Lancaster, ON He40Mh2oA wloMrlOPw 13.80 + 100 Private ownership. Managed by 45°1O'N 100hlO (Be, Ob, By, ha years Domtar Inc. Woodlot was selectively 74°27'W Bd, Bn, Bf, Cw) cut in 1988. Surrounded by agriculturalland (corn, soy). Sandy loarn; slightl~ acidic soil. . MF-2 Bainsville, ON He40Mr3oMh200h1 o(A Il.30 150 years Private ownership. Managed by 45°12.01O'N w, Bd, Bf, Be, By, Cb, ha Domtar Inc. Stand was harvested 74°25.650'W Bm, Po) -40yrs ago to build barns on farm. Surrounded by agriculturalland (corn, soy). Very fine sandy loam; some silt cla~. MF-3 Bouck's Hill, He30Mh30Bd2oBeloIdlo 5.9 ha 150 years Private ownership. Managed by ON (Hb,Aw) Domtar Inc. Stand was once part of 45°00'N large maple syrup operation. 75°12'W Surrounded by agricultural lands {ha~, so}2. Cla~ loam.

* Species codes: He = eastem hemlock; Be = American beech; Mh = hard (sugar) maple; Mr = red maple; Pw = white pine; Aw = white ash; By = yellow birch; Bd = basswood;

Ob = bur oak, Bn = black oak; Rb = bittemut hickory; Bf = balsam fir; Cw = white cedar;

Cb = black cherry; Em = elm; Po = poplar; Id = ironwood; Oh = other hardwood. Subscript numbers following species code indicate percent oftotal tree species composition occupied by that species in the stand.

100 Table 3.2. Raw species richness (Sobs), number ofindividuals, rarefaction estimated species richness (species ± SD, standardized to 120 individuals), and diversity indices of sap beetles (Coleoptera: Nitidulidae) at study sites (both trap types pooled).

Site Sobs Numberof Rarefaction Fisher's a. Simpson's individuals estimated species richness OG-1 20 310 14.87 {± 1.22} 4.77 5.28 OG-2 15 180 13.32 {± 0.58} 3.89 4.44 OG-3 15 138 14.61 {± 0.18} 4.28 5.89 MF-l 19 519 11.81 {± 1.18} 3.87 2.62 MF-2 18 646 II.07 (± 1.00} 3.43 2.28 MF-3 18 336 12.92 {± l.11} 4.06 5.10

101 Table 3.3. Indicator species analysis of sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps. Species in bold have a significant (p ~.05) Indicator Value (lndVal).

Forest Type Species Total No. Indicator p-value co Il ected value 'Old-growth' Epuraea bipunctata (n. sp.) 33.3 0.446 Epuraea erichsoni Reitter 47.0 0.423 Epuraea pe/toides Horn 16.7 1.000 Epuraea p/anu/ata Erichson 16.7 1.000 Epuraea rufa (Say) 64.2 0.420 Epuraea truncatella Mannerheim 65.5 0.144 Glischrochi/us sanguinolentus (Olivier) 53.6 0.518 Glischrochilus siepmanniW.J.Brown 33.3 0.492 Omosita discoidea (Fabricius) 43.6 0.630 Managed Brachypterus urticae (Fabricius) 16.7 1.000 Carpophi/us brachypterus (Say) 27.8 0.748 Cryptarcha ampla Erichson 16.7 1.000 Cychramus adustus Erichson 31.7 0.650 Epuraea depressa lliiger 55.3 0.582 Epuraea /abilis Erichson 62.8 0.141 Epuraea obliquus Hatch 16.7 1.000 Epuraea obtusicollis Reitter 50.0 0.192 Epuraea rufida (Melsheimer) 76.6 0.101 Glischrochilus fasciatus (Olivier) 33.3 0.471 Glischrochilus quadrisignatus (Say) 97.3 0.001 Kateretes pennatus (Murray) 16.7 1.000 Kateretes scissus Parsons 33.3 0.477 Omosita c%n (Lin ne) 52.4 0.787 Pal/odes pallidus (Palisot de Beauvois) 96.1 0.001 Phenolia grossa (Fabricius) 25.0 1.000 Sfelidofa octomaculata (Say) 8.3 1.000 Tha/ycra con c%r (LeConte) 30.6 0.471

102 Table 3.4. Sap beetles (Coleoptera: Nitidulidae) collected at study sites, 2003. Specimens in trunk window traps are in parentheses; those in flight-intercept traps are not. Totals for each species and each site are pooled numbers from both trap types.

Site S~ecies MF-1 MF-2 MF-3 Total OG-1 OG-2 OG-3 Total Brachypterus urlicae (Fabricius) 1 1 Carpophilus brachypterus (Say) 4 1 5 1 (1) 2 Carpophilus nr. rufus (female) (1) 1 Cryptarcha ampla Erichson 1 1 2 (1) 2 (1) 4 Cychramus adustus Erichson 119 119 2 4 6 Epuraea n. sp. 32 32 Epuraea nr. corlicina (female) (2) 2 Epuraea depressa lliiger 4 30 2 36 14 3 5 22 Epuraea erichsoni Reitter 2 1 3 46 1 47 Epuraea flavomaculata Maklin 1 1 Epuraea labilis Erichson 30 (1) 23 (1) 23 79 17 (1) 16 12 50 (1 ) (1 ) (3) Epuraea obliquus Hatch 1 1 Epuraea obtusicollis Reitter 4 (1) 2 3 10 (1 ) (1 ) 2 Epuraea peltoides Erichson (1) 1 1 (1) 2 Epuraea planulata Erichson 2 2 Epuraea rufa (Say) 25 23 (1) 11 60 11 53 42 106 Epuraea rufida (Melsheimer) 7 26 (1) 3 38 4 (1) 3 4 (2) 14 Epuraea truncatel/a Mannerheim (1 ) 2 1 4 6 (1) 3 (2) 2 (2) 16 Glischrochilus fasciatus (Olivier) (1 ) 2 3 Glischrochilus quadrisignatus (Say) 283(25) 364(54) 34 769 2 9 (1) 8 20 (9) Glischrochilus sanguinolentus 6 (12) 1 (1) 3 (8) 31 8 (32) 9 (5) 1 83 (Olivier) (28) Glishchrochilus siepmanni (1 ) 1 2 2 4 W.J.Brown Kateretes penna tus Murray 1 1 Kateretes scissus Parsons 13 (1) 10 24 Omosita colon (L.) 73 72 65 210 112 63 16 192 (1 ) Omosita discoidea (Fabricius) 2 5 2 9 9 4 4 17 Pal/odes pal/idus (Palisot de 22 19 33 74 2 1 3 Beauvois) Phenolia grossa (Fabricius) 1 (1) 1 1 4 1 2 3 Stelidota octomaculata (Say) 1 1 1 1 Tha/~cra concolor {LeConte~ 11 11 1 1 TOTALS 519 646 336 1501 310 180 138 628

103 --

111'--. -"", ...... -- -- 'F.-.. V_- 'Old-growtb' lAWoht Fore~t • ~~, ~ '\.

Figure 3.1. Location ofstudy sites, eastem Ontario.

104 250 A fi) 200 ii :J "0 :~ "0 150 .5 '0... Q) 100 .Q E :J Z 50

0 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ####~~~~~~~~~~~~~~~~~~ ~~~~~~~~~~~~~~p~~~~~~ ~ ,,"1 '),'>i i~' ~ ~ '),'\: '),0/ 0{]; ,,0/ '),q$ Date of Collection

25 fi) ii :J :2 20 .~ "0 .5 15 '0 ...Q) .Q E 10 :J Z 5

0 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ####~~~~~~~~~~~~~~~~~~ ~~~~~~~~~~~~~~p~~~~~~ ,,"1 ~ i~' ~ ~ '),'\: '),0/ ,,'\: ,,0/ '),q$ '\: Date of Collection

Figure 3.2. Phenology of selected Nitidulidae collected with both flight-intercept and trunk-window traps. A - Glischrochilus quadrisignatus (Say); B - Glischrochilus sanguinolentus (Olivier).

105 35 A (J'A 30 1 1 - -0 - Cychramus adustus III 25 iii :1 Pal/odes pallidus "C • 20 ~ 1 ô 0 ..CI) 15 .a 1 E :1 1 Z 10 1 5

90 80 .!!! 70 l'CI -.-Omosita colon "C= 60 :~ "C..:... 50 0 40 ~ .cCI) E 30 z= 20 10 0 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~##~////~~~~~~~~~~###~ '<:1'\ ,"~ fI," ~ ,~ ,," ,,'0 ~ .....'} 'l:fC?> "q;, fI,~ '!:>'>:> 'dq;, ,"~ fI,'>:> !V !!> ",>:> .(\ ~ ~ ,,"j '}<::f rV "j ,,<::f .(\' ~ .!$f tO' '}QI ,,'li "QI '}'<:I 'li Date of Collection

Figure 3.3. Phenology of selected Nitidulidae collected with both flight-intercept and trunk-window traps. A - Cychramus adustus Erichson and Pal/odes pallidus (Palisot de Beauvois); B - Omosita colon (L.).

106 45 40 A ~ -0-Epuraea rufa 35 1\ .R 1-- UI - .. - Epuraea labilis ii ::s \1\ 't:I 30 :~ 't:I 25 \ .5 '0 ~ \ ... 20 \ CI) 1 .Q \ E 15 0 \ ::s \ z .- 10 /\ .... )...... / -- ..... ~ /~ .~ \.~\ '-....J 5 - ... • 0"" ""J/ '-0. -0""~ "4t- \ ...... 0. J'\. 0 --~ -

20 q 18 \ - () - Epuraea erichsoni 16 • Epuraea depressa .!!! ::s111 14 +----e------.-.....:,------..•.. Epuraea rufida 't:I .. ~ 12 .5 10 '0... .! 8 E 6 z::s 4 2 0

Figure 3.4. Phenology of the most abundant species of Epuraea collected with both flight-intercept and trunk-window traps. A - Epuraea rufa and E. labilis; B - E. erichsoni, E. depressa, and E. rufida.

107 25 ~------,

1/) 20 1/) G) c .c u ii2 15 1/) ·uG) G) Co ___ MF-1 tn "'0 10 _MF-2 ---.-MF-3 ~G) Co --e-OG-1 >< W 5 ~OG-2 -z!r-OG-3

o +------~------~------~------~------~------~------~ o 100 200 300 400 500 600 700 Num ber of Individuals

Figure 3.5a. Rarefaction estimates of expected species richness (± 1 SD) of sap beetles (Coleoptera: Nitidulidae) collected with flight-intercept and trunk-window traps in six forest sites.

108 30

1/) 1/) 25 CI) c .c u 20 ii2 1/) CI) '(j 15 CI) C. tn "0 10 .! u -+- Managed forest CI) c. 5 --e-'Old-growth' forest w>< 0 0 200 400 600 800 1000 1200 Number of Individuals

Figure 3.5b. Rarefaction estimates of expected species richness (± 1 SD) of sap beetles (Coleoptera: Nitidulidae) collected with flight-intercept and trunk-window traps in managed versus 'old-growth' forests

109 Percent Similarity

100 75 50 25 o

MF-1a 1 1 MF-2a 1 MF-1b 1 MF-2b MF-3a MF-3b OG-1a OG-2a 1 OG-1b J OG-2b OG-3a 1 OG-3b 1

Figure 3.6. Dendrogram of c1uster analysis on species of sap beetles (Coleoptera: Nitidulidae) collected from flight-intercept traps.

110 Appendix 3.1a. Sap beetles (Coleoptera: Nitidulidae) coUected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #1, Cornwall, ON, (OG-l).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S~ecies Ma:l Ma:l Ma:l Ma:l Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Se~ Se~ Se~ Se~ Oct Brachypterus urticae (Fabricius) CarpophiJus brachypterus (Say)

Carpophilus nr. rufus (female) (1 ) Cryptarcha ampla Erichson (1) Cychramus adustus Erichson 1 Epuraea n. sp. 8 6 18 Epuraea nr. corticina (female)

Epuraea depressa lliiger 3 2 1 3 2 2 Epuraea erichsoni Reitter 3 11 19 11 - Epuraea flavomaculata Maklin - Epuraea labilis Erichson 2 3 2 3 (1 ) 2 2 Epuraea obliquus Hatch

Epuraea obtusicollis Reitter (1 ) Epuraea peltoides Erichson 1 Epuraea planulata Erichson 2 Epuraea rufa (Say) 3 (1) 3 4 Epuraea rufida (Melsheimer) 2 Epuraea truncatella Mannerheim 1 (1) 2 Glischrochilus fasciatus (Olivier) Glischrochilus quadrisignatus (Say) Glischrochilus sanguinolentus (Olivier) (6) (3) 1(10) (1 ) (1 ) 1(1) 2(4) 2(4) (1) (1) GlishchrochiJus siepmanni W.J.Brown 2 Appendix 3.1 a. continued

Omosita c%n (Linné) 5 (1) 11 3 7 3 34 26 11 4 4 Omosita discoidea (Fabricius) 2 6 Pal/odes pallidus (Palisot de Beauvois) Phenolia grossa (Fabricius) Stelidota octomacu/ata (Say) Tha/ycra conc%r (LeConte)

...... N Appendix 3.lb. Sap beetles (Coleoptera: Nitidulidae) coUected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #2, Lancaster, ON, (OG-2).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Brachypterus urticae (Fabricius) Carpophilus brachypterus (Say) 1(1) Carpophilus nr. rufus (female) Cryptarcha ampla Erichson Cychramus adustus Erichson Epuraea n. sp. Epuraea nr. corticina (female) Epuraea depressa IlIiger 1 Epuraea erichsoni Reitter - Epuraea flavomaculata Maklin -w Epuraea labilis Erichson 2 3 3 (1) 2 3 Epuraea obliquus Hatch Epuraea obtusicollis Reitter Epuraea peltoides Erichson Epuraea planulata Erichson

Epuraea rufa (Say) 7 14 21 5 3 Epuraea rufida (Melsheimer) 2 Epuraea truncatella Mannerheim 1(2) Glischrochilus fasciatus (Olivier) Glischrochilus quadrisignatus (Say) 2 (1 ) 3 Glischrochilus sanguinolentus (Olivier) 4 3(1) (4) Glishchrochilus siepmanni W.J.Brown 2 Appendix 3.lb. continued

Omosita c%n (Linné) 7 6 7 5 7 6 3 7 2 2 4 2 Omosita discoidea (Fabricius) 1 Pal/odes pallidus (Palisot de Beauvois) Phenolia grossa (Fabricius) Ste/idota octomacu/ata (Say) Tha/ycra conc%r (LeConte)

...... ~ Appendix 3.1c. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in old-growth forest #3, Morrisburg (Upper Canada), ON, (OG-3).

Cryptarcha ampla Erichson (1) Cychramus adustus Erichson 2 Epuraea n. sp. Epuraea nr. corticina (female) Epuraea depressa lliiger 2 2

...... Epuraea erichsoni Reitter ...... Vl Epuraea flavomaculata Maklin (1) Epuraea labilis Erichson 2 2 2 2 Epuraea obliquus Hatch Epuraea obtusicollis Reitter

Epuraea peltoides Erichson (1) Epuraea planulata Erichson Epuraea rufa (Say) 9 3 3 9 11 5 2 Epuraea rufida (Melsheimer) 2 1 Epuraea truncatella Mannerheim (2) Glischrochilus fasciatus (Olivier) Glischrochilus quadrisignatus (Say) Glischrochilus sanguinolentus (Olivier) 1(5) (3) (2) (6) (3) (4) (1) (1) (1) (2) Glishchrochilus siepmanni W.J.Brown Appendix 3.Ie. eontinued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Kateretes penna tus (Murray) Kateretes scissus Parsons Omosita c%n (Linné) 2 5 4 2 Omosita discoidea (Fabricius) 2 Pal/odes pallidus (Palisot de Beauvois) Phenolia grossa (Fabricius) 2 Stelidota octomacu/ata (Say) Tha/ycra conc%r (LeConte)

...... 0\ Appendix 3.1d. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #1, Lancaster, ON, (MF-l).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Brachypterus urticae (Fabricius) Carpophilus brachypterus (Say) Carpophilus nr. rufus (female) Cryptarcha ampla Erichson Cychramus adustus Erichson Epuraea n. sp. Epuraea nr. corticina (female) Epuraea depressa IIliger Epuraea erichsoni Reitter - Epuraea flavomaculata Maklin --....1 Epuraea labilis Erichson 7 8 2 2 2 3 1(1) Epuraea obliquus Hatch

Epuraea obtusicollis Reitter (1) 2 Epuraea peltoides Erichson (1 ) Epuraea planulata Erichson Epuraea rufa (Say) 8 5 4 2 Epuraea rufida (Melsheimer) 3 Epuraea truncatella Mannerheim (1 )

Glischrochilus fasciatus (Olivier) (1 ) Glischrochilus quadrisignatus 120(2 (Say) ) 6 57 21 4(2) 3 3 2 3 19(1) 8(20) 3 2 3 8 18 Glischrochilus sanguinolentus (Olivier) 3(6) (2) 1(2) (1) (1) Glishchrochilus siepmanni W.J.Brown (1) Appendix 3.1d. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Kateretes pennatus (Murray) Kateretes scissus Parsons 2 3 2(1) 2 3 Omosita c%n (Linné) 23 10 5 1 8 2 11 3 3 Omosita discoidea (Fabricius) 2 Pal/odes pallidus (Palisot de Beauvois) 2 3 5 2 4 Pheno/ia grossa (Fabricius) (1 ) Stelidota octomacu/ata (Say) Tha/ycra conc%r (LeConte)

...... 00 Appendix 3.1e. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #2, Bainsville, ON, (MF-2).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Brachypterus Ulticae (Fabricius) 1 Carpophilus brachypterus (Say) 1 Carpophilus nr. rufus (female) Cryptarcha amp/a Erichson Cychramus adustus Erichson Epuraea n. sp.

Epuraea nr. corticina (female) (2) Epuraea depressa IlIiger 4 4 3 2 3 8 5 Epuraea erichsoni Reitter ...... \0 Epuraea flavomacu/ata Maklin Epuraea /abilis Erichson 5(1) 2 2 1(1) 2 3 2 4 Epuraea obliquus Hatch Epuraea obtusicollis Reitter Epuraea pe/toides Erichson Epuraea p/anu/ata Erichson

Epuraea rufa (Say) 1 2 2 7 3(1) 2 3 Epuraea rufida (Melsheimer) 9 2 (1) 3 2 2 2 4 Epuraea truncatella Mannerheim 2 Glischrochilus fasciatus (Olivier) Glischrochilus quadrisignatus 31(16 (Say) 74(3) 2(1) 37(1) 2 5(9) (2) 4(8) 3(5) 6 7(8) ) 17(1) 16 18 5 4 13 70 16 34 Glischrochilus sanguino/entus (Olivier) (1 ) Glishchrochilus siepmanni W.J.Brown Appendix 3.1 e. continued

7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Kateretes penna tus (Murray) Kateretes scissus Parsons 2 2 2 1 2 Omosita c%n (Linné) 15 2 8 7 7 16 2 2 2 3 3 Omosita discoidea (Fabricius) 3 2 Pal/odes pa/lidus (Palisot de Beauvois) 3 2 6 2 2 1 Pheno/ia grossa (Fabricius) Stelidota octomacu/ata (Say) Tha/ycra conc%r (LeConte)

...... N o Appendix 3.1f. Sap beetles (Coleoptera: Nitidulidae) collected in flight-intercept traps (no parentheses) and trunk-window traps (in parentheses) in managed forest #3, Bouck's Hill (near Williamsburg), ON, (MF-3).

Date of collection 7 14 21 28 4 11 18 25 2 9 16 23 30 6 13 20 27 3 10 17 24 3 S~ecies Ma:z:: Ma:z:: Ma:z:: Jun Ma:z:: Jun Jun Jun Jul Jul Jul Jul Jul AU!i! AU!i! AU!i! AU!i! Se~ Se~Se~ Se~ Oct Brachypterus urlicae (Fabricius) Carpophilus brachypterus (Say) Carpophilus nr. rufus (female) Cryptarcha ampla Erichson

Cychramus adustus Erichson 3 16 30 30 10 8 10 4 5 2 1 Epuraea n. sp. Epuraea nr. corticina (female)

Epuraea depressa lliiger 1 Epuraea erichsoni Reitter ..... N ..... Epuraea flavomaculata Maklin Epuraea labilis Erichson 3 1(1) 1(1 ) 6(1) 2 3 2 Epuraea obliquus Hatch

Epuraea obtusicollis Reitter 3 Epuraea peltoides Erichson Epuraea planulata Erichson

Epuraea rufa (Say) 3 5 2 Epuraea rufida (Melsheimer) 2 Epuraea truncatella Mannerheim Glischrochi/us fasciatus (Olivier) Glischrochilus quadrisignatus (Say) 2 1(1) 2 5(5) 7 (1) 7(2) 3 4 Glischrochilus sanguinolentus (Olivier) (4) (1 ) (1 ) (2) Glishchrochi/us siepmanni W.J.Brown Appendix 3.1 f. continued

7 14 21 28 4 11 18 25 2 9 1623 30 6 13 20 27 3 10 17 24 3 Species May May May May Jun Jun Jun Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Sep Sep Oct Kateretes pennatus (Murray) Kateretes sCÎssus Parsons

Omosita c%n (Linné) 14 14 12 7 2 5 5 2 Omosita discoidea (Fabricius) 1 1 Pal/odes paJ/idus (Palisot de Beauvois) 2 3 4 4 6 2 5 2 4 Pheno/ia grossa (Fabricius) Stelidota octomacu/ata (Say) Thalycra conc%r (LeConte) 2 3 2 1

IV -IV CHAPTER 4: General Conclusion This study has contributed valuable data on the diversity and distribution patterns ofnumerous species ofsaproxylic beetles (Coleoptera) in the forests ofsoutheastern Ontario, Canada. Specifically it has provided infonnation on several families of cryptic, often overlooked beetles, inc1uding members of the Leiodidae and Nitidulidae. While lengthy studies on saproxylic Coleoptera abound in Fennoscandia, this was the first season-long biodiversity study in eastern Ontario that focused specifically on fungivorous Coleoptera. Likewise, very few studies focus on nitidulid diversity, and this study was one of the first to provide an unbiased survey of the nitidulid species present in small hemlock-hardwood forest stands in North America. Several of the forest sites are designated as "Demonstration Forests" by the Eastern Ontario Model Forest and are used to educate landowners on ecologically sound forest management procedures. In addition to contributing to our knowledge of the distribution ranges ofmany species, this study has also provided baseline infonnation for future monitoring, research and/or educational work in the area's forests. By comparing managed to unmanaged 'old-growth' forests, this study has provided valuable infonnation to forest managers and conservation workers on the impact that forest harvest operations have on a valuable component of the forest ecosystem. Round fungus beetles (Leiodidae: Leiodinae) were the most frequently sampled fungivorous beetles, with 87% of all species collected belonging to this family. Leiodids are small beetles specializing on a variety of slime mold and fungus species in forested habitats. Flight-intercept traps are ideal for collecting these small beetles. Contrary to predictions, the species richness and abundance of fungivorous Coleoptera was similar in both managed and 'old-growth' forests. Several other published studies have found significant differences in fungivorous and/or saproxylic fauna between managed and unmanaged habitats. There are several possible explanations for the result ofthis study, inc1uding: beetles chosen for the study are not vulnerable to the habitat alteration caused by forest harvest operations occurring in southeastern Ontario, harvest may not yet have had time to affect the beetle assemblages, effects ofharvest may have already been mitigated by time, specialist beetles may have long ago disappeared from the sites due to

123 extensive early fragmentation, or managing cut stands for 'old-growth' characteristics may provide sufficient habitat (dead wood) for these beetle species. Unlike the beetles studied in chapter 2, the species abundance of nitidulid beetles (chapter 3) was higher in managed forests and the species richness was higher in 'old growth' forests. However, much ofthis result can be attributed to the dominance of Glischrochilus quadrisignatus in several of the sites. G. quadrisignatus is a common nitidulid and is known to be attracted to maize. The presence of corn fields surrounding two of the managed sites likely acted as a 'bait' to this beetle, greatly increasing its abundance and skewing the overall diversity and abundance results of managed forests. Examining the seasonal abundance patterns of the most commonly collected nitidulid beetles revealed distinct abundance peaks for several species. Most important was the seasonal division in the abundance of Pal/odes pallidus and Cychramus adustus, both species that specialize on fleshy white fungus. Several species of fungivorous beetles showed strong or significant associations with either managed or 'old-growth' forests. Triplax macra was strongly associated with 'old-growth' forests, while Anisotoma blanchardi, Anogdus obsoletus, and Mycetina perpulcra were significantly associated with managed forests. The nitidulid Pal/odes pal/idus was strongly associated with managed forest while Glischrochilus sanguinolentus, a nitidulid that was more frequently collected with trunk-window traps than with flight-intercept traps, may have had a possible association with 'old-growth' forest. Anisotoma inops, a species previously considered as a potential 'indicator' of 'old-growth' forests in north-eastern North America, was found in both managed and 'old-growth' forests in this study and thus may be associated with mature forests, managed or not. Moreover, Anisotoma inops was collected far more often in trunk window traps and this may have influenced the conclusions made by other studies which only used large-area flight-intercept traps to capture beetles. Future work in this area should include more detailed inventory of the downed woody debris in forests. AIso, a more comprehensive survey of the fungi present in each stand would improve forest comparisons and conclusions based on species specializations. Comparative arthropod biodiversity studies are important and deserve our continued attention. Arthropods comprise almost 64% of global biodiversity and

124 forests harbour a diverse array ofthese organisms. With continued use of our forest resources, it is imperative that we understand the effect that harvest and management practices have on aIl components ofthe forest ecosystem, especially on smaller organisms often overlooked in management plans and large-scale studies.

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