UTILITY OF GROUND BEETLES (COLEOPTERA:

CARABIDAE) AS INDICATOR SPECIES FOR MONITORING

BIODIVERSITY EFFECTS FROM VARIABLE RETENTION

HARVESTING PRACTICES

BC FOREST SCIENCE PROGRAM

PROJECT NUMBER: Y061029

2005/06 FINAL REPORT January 2006

BY Isobel A. Pearsall

Pearsall Ecological Consulting, 99 Machleary Street, Nanaimo, B.C. V9R 2G3 Email: [email protected] Contract report to: W.J. Beese, Cascadia Forest Products 4th Floor, 65 Front Street, Nanaimo, BC V9R 5H9

TABLE OF CONTENTS

Table of Contents ...... 2 Executive Summary...... 3 Introduction...... 5 Objectives...... 7 Materials and Methods...... 7 Study sites ...... 7 Method of trapping ...... 10 Sampling design...... 10 Data collection ...... 13 Data Analysis...... 13 Trap catches ...... 13 Description of carabid species ...... 13 Patterns of abundance ...... 13 Statistical analysis...... 14 By-Catch data...... 16 Results ...... 17 Ground vegetation and stand differences among sites...... 17 Trap catches ...... 17 Description of carabid species ...... 17 Patterns of abundance ...... 30 Statistical analysis...... 41 Discussion...... 77 Conclusions...... 86 Acknowledgements ...... 87 References...... 88 Appendix One: Site Characteristics 2005...... 92 Appendix Two: Photographs of Sites...... 94 Appendix Three: New species discovered during 2005 ...... 108 Appendix Four: Analysis of by-catch...... 109

2 EXECUTIVE SUMMARY

• During 2005, I began a detailed examination of the effect of patch size on carabid communities in both experimental and operational forest blocks in West Island Timberlands 2nd growth and old growth. In particular, I was interested in comparing how well patches of different size retain original forest communities of beetles, as well as to examine edge effects, the potential “lifeboating” of patches, and how the size of patches affects composition of beetle communities within the cut matrices. I was also interested in determining whether the results were consistent among old growth and 2nd growth sites.

• A total of three group retention sites (G1, G2, G3) were chosen in 2nd growth forest, and two group retention sites (G4 & G5) were chosen in old growth forest in West Island Timberlands. Site G1 contained large patches whereas those in site G2 were small. Both small and large patches were found in site G3. Site G4 had small patches and site G5 had two large patches. Traps were also set in 2nd growth forest located adjacent to site G2 as our 2nd growth control site (S1). We also set traps in the Klanawa group size VRAM site, which consisted of 5 treatments: old growth control (O), small patches (Sm), medium size patches (M), large patches (L) and clearcut ( C). All four 2nd growth sites were in the CWHxm and all old growth sites were in the CWHvm1. All sites had been harvested in 2003/2004 except for the VRAM site which was harvested in 2005. All 3 VRAM treatments (small patches, medium patches and large patches) had the same retention level of 24.3%. All the other sites had similar retention levels around 18-25% with the exception of site G5, where retention levels were 39.9%.

• 3 years post-harvest, the patches were able to retain forest specialists in all of the group retention sites. Carabid beetles showed a response to patch size in group retention sites, with higher catches per trap of forest specialists in larger patches than in smaller patches. This response was seen in both old growth and 2nd growth sites. I have been able to produce species-patch area curves for the most abundant species, showing a clear response in terms of increased abundance to increasing patch size by forest specialists. However, retention of forest specialists was generally better in old growth patches than 2nd growth patches, and small patches of 2nd growth sites were particularly poor at retaining species such as S. angusticollis, Z. matthewsii, and P. crenicollis.

• Carabid beetles showed edge responses, and several species built up at or close to the edge of patches. This may be associated with a general unwillingness by forest specialists to venture further out into the matrix. However, some of the generalist species also built up at the edges, which suggests that this may be related to the change in vegetation or microclimate at this location.

3 • The 2nd growth sites had generally greater species richness and diversity than the old growth sites, and control sites in both forest types generally had lower species richness and diversity than the group retention sites, probably because of influx of new species associated with new habitats and forest edges that were provided by group retention harvesting. Diversity was generally greater in small patch sites than sites with larger patches, except in the VRAM site, where diversity indices were very similar among the group retention treatments. Higher diversity in the small patch sites may be associated with the greater amount of edge, and greater size of matrix tracts, which may influence forest specialists in a negative fashion, and allow for greater abundance of forest generalists.

• Seasonal patterns of abundance showed general peaks of abundance in May and September in most sites, for both total carabids pooled and for Scaphinotus angusticollis, the most commonly captured beetle. These months coincide with reproductive periods, and generally, both activity and abundance are greatest at this time.

• Our results show a strong response by carabid beetles to forest edges and patch sizes. Whether this is a result of the microclimate or prey base of small patches being less ideal than larger patches, or an isolation effect, remains to be determined. Another factor that may be important to consider is the effect of patch shape, as my study only examined the factors of patch size and age. Further work in Port McNeil and in West Vancouver Island over 2006 will allow for a further opportunity to examine these responses in different variants.

4 INTRODUCTION

The adaptive management program of Cascadia Forest Products (formerly Weyerhaeuser) was designed to examine the effectiveness of variable retention systems and stewardship zoning in maintaining forest attributes that may be necessary for sustaining biodiversity and ecological functions. To aid in this process, it has been necessary to determine whether there may be indicator organisms that may be useful to assess ecosystem health. Such indicator organisms would ideally be individual species, groups of species or structures that perform critical ecosystem functions or are particularly sensitive to disturbance. Specific details of the utility of carabids in this regard were outlined in the initial project proposal (Pearsall 2000). During the first year of this pilot study, 2001, work was carried out in Northern Vancouver Island to gather baseline information about carabid species abundance and distribution in mature, immature and clearcut forests at low and high elevations. A secondary objective was to evaluate costs and sampling efficiency to determine the suitability of ground beetles as focal species for the adaptive management program. During 2002, work was continued in two experimental VR-sites to assess how carabid populations responded to two methods of variable retention forestry: dispersed retention and group % retention. During 2003, work was continued in operational sites in Douglas-fir forests of South Vancouver Island and Western Hemlock forests in North Vancouver Island. Factors of interest were fourfold. Firstly, work was done in a number of different ages of stands to fill in gaps of ages done during 2001. Other issues of importance were a detailed examination of edge effects, a retrospective examination of possible effect of changing matrix age/lifeboating potential of remnants, and an examination of remnants and extrapolation to possible changes within retention patches over time. During 2004, we examined the oldest operational VR that was available in West Vancouver Island and examined the communities of carabids among group and dispersed retention sites with differing levels of tree retention. We also sampled a 2nd -growth control site in the same region, which allowed us to determine how well the original communities of carabids were able to persist in the operational VR sites.

5 Fragmentation and habitat loss are among the most important causes worldwide of species decline (Haila et al. 1994; Didham et al. 1996). As habitat is lost, consequences of fragmentation include decreased fragment size and increasing amounts of edge (Harris 1984, Matlack & Litvaitis 1999). Small forest fragments with a high proportion of edge habitat are vulnerable to invasion by species from the surrounding cut matrices. Edge habitat is unsuitable for some species that require interior forest habitats for survival, and thus such species may be lost if fragments become too small (Stevens & Husband 1998, Haila 1999). It is generally understood that there may be threshold patch sizes, below which immigration and reproduction would be too low to maintain the patches as adequate source areas for certain species, including forest carabid beetles. Overall, the studies that I have carried out over the past 4 years have established the change in carabid communities with forest age, and I have begun to develop a VR type and level response curve. However, I have not yet established a response curve for carabid beetles to the size of retained patches. It has been suggested that forest managers should minimize adverse edge effects by leaving fragments that are large enough to maintain interior forest specialists (e.g. Spence et al. 1996; Burke and Goulet 1998; von Sacken 1998). Results of my work done in experimental group VR blocks on North Vancouver Island during 2002 indicated that patches are able to retain the original old growth forest communities of beetles, at least in the short term. Increasing the level of tree retention resulted in improved retention of forest beetle species. However, patches in 30% retention treatments were generally larger than patches in 20% or 10% retention, and thus % tree retention, size of patches and proximity of patches were confounded. Recent work in West Vancouver Island during 2004 showed that operationally harvested VR patches were unable to retain the original forest species. I suggest that this is related to the size of the patches, as the operational patches were generally much smaller than those set up in the experimental sites. During 2005, I began a detailed examination of the effect of patch size on carabid communities in both experimental and operational forest blocks in West Island Timberlands. In particular, I was interested in comparing how well patches of different size retain original forest communities of beetles, as well as to examine edge effects, and how size of patch affects composition of beetle communities within the cut matrices.

6

OBJECTIVES

The main questions addressed during the 2005 field season were:

1) What are the differences in carabid beetle assemblages found in the large and small group VR sites sampled in West Vancouver Island? 2) Are there differences among the 2nd-growth control and the patches in small and large patches of 2nd growth group retention cutblocks in terms of beetle assemblages, diversity and abundance? Are there differences among the old growth control and the patches in small, medium and large patches of old growth group retention cutblocks in terms of beetle assemblages, diversity and abundance? In both 2nd and old growth forests, have patches been able to conserve the original carabid communities associated with the un-cut forest? How do the communities of the relevant control sites and the group retention cutblocks vary overall, when matrix and patch communities are taken into account? 3) What are the differences in carabid catches between the matrices of the different group retention blocks? 4) Are there any edge effects? 5) Are large patches better than small patches at conservation of forest specialists, and are these differences consistent among old growth and 2nd growth sites?

MATERIALS AND METHODS

Study sites

A total of three group retention sites (G1, G2, G3) were chosen in 2nd growth forest, and two group retention sites (G4 & G5) were chosen in old growth forest in West Island Timberlands. Site G1 contained large patches (approximately 80 m by 80 m), whereas those in site G2 were small (approximately 50 m by 50 m). Both small and large patches were found in site G3 (50 m by 50 m for the small, and approximately 200 m by 50 m for the large). Site G4 had small patches (50 m by 50 m) and site G5 had two large patches (300 m by 100 m). Traps were also set in 2nd growth forest located adjacent to site G2 as our 2nd growth control site (S1). We also set traps in the Klanawa group size

7 VRAM site, which consisted of 5 treatments: old growth control (O), small patches (Sm), medium size patches (M), large patches (L) and clearcut ( C). The large patches were approximately 100 m by 100 m, the medium sized patches were 60 m by 60 m and the small patches were approximately 27 m by 25 m. All four 2nd growth sites were in the CWHxm and all old growth sites were in the CWHvm1. All sites had been harvested in 2003/2004 except for the VRAM site which was harvested in 2005. All 3 VRAM treatments (small patches, medium patches and large patches) had the same retention level of 24.3%. All the other sites had similar retention levels around 18-25% with the exception of site G5, where retention levels were 39.9%.

8 Table 1. Characteristics of sites sampled during 2005.

Block Harvest Age of site Species Name Block Complete Planted % Retention Variant Elevation composition 2nd growth 61 17.8 CWHxm (vm1 280-400m FdCwHwDr/Mb G1 973405 April 2003 March 2003 influence) Winter 62 26.3 CWHxm 100 to 250 Fd45% Hw40% 2003/Spring September 2004 and meters Cw10% DrMb5% G2 062204 2004 March 2005 62 20.8 CWHxm 200-400m Fd80% Hw10% G3 964402 Spring 2004 March 2005 Cw5% DrMb5 62 100 CWHxm 200-400m Fd80% Hw10% S1 N/A Cw5% DrMb5 Old growth Old March 2005 18.4 CWHvm1 240 to 560 m CwHwBa(Fd) G4 862118 2003/2004 growth Old 39.9 CWHvm1 380 to 650 m HwCwBa G5 861418 May 2004 growth September 2005 Old Not planted. 24.3 CWHvm1 120 to 600 m CwHwBa 2005 growth Scheduled for VRAM 86622 March 2006 Tree species: Fd (Douglas-fir), Cw (Western red-cedar), Hw (Western Hemlock), Ba (Balsam fir), DrMb (Alder, Bigleaf Maple)

A total of 291 traps were placed out in the operational (G1-G5, S1) and experimental sites (old growth and medium sized patches) in late April, and a further 86 were placed out in the experimental site (small patches, clearcut and large patch treatments) in late May. It was not possible to get access to the small patch, clearcut or large patch treatments until later in May and thus, these sites were set up one month later than the other sites. Collections were made every month for May-September inclusive.

Method of trapping

As in 2001 -2004, pitfall trapping was used to trap invertebrates. The design of these traps may be found in Pearsall (2002). The contents of each pitfall trap were placed into labelled collecting vials, which were filled with isopropyl alcohol back at the lab. The vial contents were analysed prior to the following field trip.

Sampling design

In all the group retention sites, traps were placed within patches, at the patch edge, and within the matrices of the cut blocks. In each site, 3 transects were set up radiating from each of 3 patches. Each transect was 1200 apart from the next, with the first cardinal direction determined randomly. For sites with either small or medium sized- patches, 12 traps were placed in each patch, with 3 inside the patch, 3 at the edge, 3 in the matrix 5-10 m from the edge, and a final 3 located 25 m further into the cut matrix (see Figure 1 for idealized layout in this treatment type). For sites with large patches, 16 traps were placed in each patch, with 3 inside the patch, 3 5-10 m within the patch from the patch edge, 3 at the edge, 3 in the matrix 5-10 m from the edge, and a final 3 located 25 m further into the cut matrix (see Figure 2 for idealized layout in this treatment type). In the clearcut, old growth and second growth sites, we positioned trapping stations along three systematically chosen transect lines. The direction of transects were altered as required if they coincided with features such as cliff faces, streams, or if the trap would be located too close to the edge i.e. less than 50 m from the edge of the block. Mapping of sample stations allowed for easy relocation of traps. A total of 15 pitfall traps (were

10 dug into the clearcut and old growth treatments, and 12 traps were placed in the 2nd growth control site (see Figure 3 for idealized layout of these traps). Each trap was placed at least 25 m apart.

11 Figure 1. Idealized example of the layout of pitfall traps in the clearcut, old growth and 2nd growth control sites

Old growth site

50M FROM Trapping EDGE Station

Traps shown above were placed every 25 m along transect lines, and the first station was >50 m into the block from the road.

Figure 2. Idealized example of layout of traps radiating from small patches in the group retention sites (large patches had one extra trap 5-10m into the block from the block edge..

Cut matrix Transect 1

Patch

Transect 2 Transect 3

12

Data collection

To try to examine between-site differences, a simple categorization of ground vegetation was made in each block. Photographs of each site were taken over the year. Some habitat and vegetation information for the experimental VRAM site was available from the Access database VR-mdb provided by Jeff Sandford, Weyerhaeuser Nanaimo. This database housed the information collected by the structural monitoring group pre- harvest in two of the dispersed retention sites, specifically: % ground coverage by trees, shrubs, mosses and herbaceous vegetation, height of shrubs and herbs, the dominant 3 herb and shrub species, litter characteristics, course woody debris, and dominant tree species as well as other details regarding decay and disease classes for trees. Numbers and species of carabid (ground) beetles were noted for every trap for all months. All other beetle groups were also identified to the family level and counted for all visits. Carabid and other beetles were pinned where necessary and taken to the George J. Spencer Entomological museum for aid with identification. All other by-catch also was assessed for all sampling dates, although many of these groups were only identified to Order.

DATA ANALYSIS

Trap catches

All carabid data were subjected to the following analyses:

Description of carabid species

The abundance and types of species collected in each block were listed, and any relevant information regarding these species’ biology was noted in an appendix.

Patterns of abundance

We examined patterns of abundance of the most commonly captured species.

13

Statistical analysis

Individual species differences among treatments and locations within group retention types. This year, I was primarily interested in comparing how well original carabid communities were retained in the differently sized patches. I was not specifically interested in the effects of retention level during this study, but rather the effect of patch size. Firstly, I used paired t-tests to compare catches of the most commonly captured forest species in patches of each site with the relevant matrix site catches. This was done to see whether the patches in the different sites were equally able to retain the different forest specialist beetle species. Next, t-tests were used to compare the catches of forest specialists in the matrices of the old growth group sites with catches in the clearcut treatment of the VRAM experimental site. This was done to allow me to examine whether carabids, in particular, forest specialist species, were more abundant in the matrices of group VR patches than in the clearcut, and whether this varied with patch size. This comparison was done only for old growth sites, as we did not have a clearcut treatment in the 2nd growths sites. Next, t-tests were used to compare the following: a) catches within large patches and small patches in site G3, b) catches from small patches of site G2 with large patches of site G1, c) catches in small patches of site G4 with large patches of site G5 and d) to compare the catches among small, medium and large patches of the VRAM site. I focussed on comparisons of the forest specialist species, and this allowed me to determine whether large patches were better able to retain forest specialist species than small patches. Next, one-way ANOVA was used to compare the individual species abundance among the group retention patch catches (of 2nd growth sites) and the 2nd growth control, and among the group patch catches (of old growth sites) and the old growth control, to determine whether the patches were able to maintain similar communities to the original forest.

14 Next, a General Linear Model procedure (Zar 1984) was used to compare the responses of individual species to treatments within patch retention cuts. Comparisons were done among old growth and 2nd growth sites separately. I focused on the most commonly captured species, including both forest specialist species and generalist species, and examined the effects of site (block ie G1, G2, G3 etc), location (In, Out, Edge, 5-10m out from edge, 5-10m in from edge) and location nested within sites. Finally, the response of the most abundant species to patch size was assessed by plotting species-area curves, where average patch size from each site was estimated from setting maps

Species evenness. Whittaker plots (Krebs 1989) were produced whereby the log of % abundance for each species was plotted against the species rank for each site, and, for group retention sites, for each trap location (In, Out, Edge, 5 to 10m out from Edge, 5 to 10m in from Edge). These plots allow a simple visual comparison of the dominance or equality of species within the community. Total numbers of carabids caught in each particular treatment were summed for the whole season and the sum of catches of each species were then expressed as a percentage of this yearly total.

Heterogeneity measures. Several nonparametric diversity indices were calculated for each treatment type (Southwood 1978). These measures take into account both species richness and species evenness and are thus referred to as heterogeneous measures (Magurran 1988, Krebs 1989). They are valuable in that they allow for comparison across all treatments, despite differences in sample size and number of individuals caught, and are useful as they assume no statistical distribution. The Simpson index, 1-D and the Shannon Wiener index, H’ are useful values in studies of this kind. These indices combine the effects of species richness and dominance within a community and allow for comparison across treatments. The Simpson index highlights the changes that occur in the most common species and the Shannon-Wiener

15 index, a measure of entropy within the system, highlights the more rare species in the assemblage (Magurran 1988). Using the EstimateS software (Colwell, 2004), a number of other indices were calculated, including the alpha mean, which is Fisher's alpha diversity index, and two different estimators of expected species richness, the ACE mean and the ICE mean. These indices were calculated for the carabid beetle data only.

Distance coefficients. To assess how the different sites compare to one another, a distance matrix was calculated using Euclidean distance metric. This allows us to visualize the dissimilarity of the different communities sampled from the different block types. The distance matrix was imported into a simple linkage cluster analysis program (in Systat) to produce a tree diagram. This was done both for catches by site and also for catches by site and by location of traps (ie. for group retention sites, for each trap location (In, Out, Edge, 5 to 10m in from Edge, 5 to 10m out from Edge)). For the group sites, I used weighted data for total site catches, such that abundance of carabids from traps in any particular location were weighted by the estimated area made up by that particular location within a block, to allow for a more realistic estimate of community composition.

By-Catch data

The data for staphylinid beetles, curculionid beetles (weevils), snails, crickets and spiders were extensive enough that some analyses could be performed. Examination of temporal trends were made for these groups. The numbers for other by-catch, including shrews and salamanders, were low at both sites for most dates, and were not examined. All by-catch analyses, results and discussion are given in Appendix Four, together with Tasheena Gardner‘s honours thesis paper from SFU in which she examined the millipede by-catch caught during 2005.

16

RESULTS

Ground vegetation and stand differences among sites

The second growth sites were all located in the CWHxm and were dominated by Douglas-fir (Pseudotsuga menziesii) whereas the 2nd growth sites were located in CWHvm1 and were dominated by western hemlock (Tsuga heterophylla) and Western Red-cedar (Thuja plicata). In the Appendix we note the major herb and shrub species that we found. Larger patches in both second growth and old growth sites had retained mosses and a rich ground vegetation, but small patches in 2nd growth sites were generally drier and more barren than was the case in small patches in old growth sites. No doubt the higher levels of moisture in the CWHvm1 were able to offset the microclimatic effects on vegetation structure in small patches, which were apparent in the 2nd growth sites in the CWHxm.

Trap catches

Catch data for all animals caught in pitfall traps was placed into an Access 2000 Database. A synoptic collection of carabid beetles has been given to the George J. Spencer Entomological Museum and one remains with me. The latter collection includes pinned examples of the non-carabid beetles captured. Beetles are identified by site, location and pitfall trap, date of collection, and species/genus name. The other by-catch has been placed in glass jars of isopropyl alcohol and pooled by site and date.

Description of carabid species

All carabid beetles were identified at the species level. Overall, there were 17 species of carabid beetles, and 9900 specimens collected over the season. Overall, fewest species of carabids (5) were collected in the clearcut treatment, and most species (between 13-14) were collected in the 2nd growth group VR sites (Table 1). There were mores species in the 2nd growth group VR sites than found in the adjacent 2nd growth control (9 species). When we compared the number of species by patch size in site G3,

17 we found 13 species in the small patches, and 14 species in the large patches. The number of species of carabids collected in the old growth sites varied between 9 and 11. I used Lindroth (1961-69), and Kavanaugh (1992) to determine what is known about each species in terms of its diet, habitat requirements, and its flight capability. For many species, records of observed flight may not have been made. In this case, the presence of fully formed wings and wing muscles may have been used to draw the conclusion that flight capabilities are complete (Lindroth 1961-69). Details about some of the new species captured this year are listed in an appendix at the end of this report (Appendix Four). Details of the other species are listed in Pearsall (2002, 2003, 2005) final reports for work carried out during 2001-2004. By site, the averages catches of carabids for all locations pooled for the entire season were greatest overall in the 2nd growth control, followed by G1, G3 Large Patches, G5, O, M, G4, G3 Small patches, G2, Sm and lowest overall in the clearcut. For the second growth sites, highest catches per trap were from the 2nd growth control, followed by G1 (large patches), G3 (large patches), G3 (small patches), and lowest in G2 (small patches). For the old growth sites, catches were similar and highest in G5 (large patches) and the old growth control, followed by M (medium patches), G4 (small patches), Sm (small patches) and lowest overall in the clearcut. In both 2nd growth and old growth sites, catches appeared to decline with reducing patch size. Overall, Scaphinotus angusticollis was the most common carabid species captured, followed by Pterostichus crenicollis, P. herculaneus, P. lama and Zacotus matthewsii. In the 2nd growth sites, the most abundant species captured varied by site. In the 2nd growth control, S. angusticollis was the most common, followed by P. herculaneus. In the large patch site G1, S. angusticollis was the most common, followed by P. crenicollis. In the small patch site G2, P. herculaneus was the most common, followed by P. algidus. In the small patches of site G3, S. angusticollis was the most common, followed by P. herculaneus. In the large patches of site G3, S. angusticollis was the most common, followed by P. herculaneus, but large numbers of P. crenicollis were also captured in this site. In all of the old growth sites, S. angusticollis was the most common, followed by P. crenicollis, with the exception of the small patch site of the

18 VRAM experiment. In that site, S. angusticollis was the most commonly captured species, but the 2nd most commonly caught species was P. herculaneus.

Overall, the largest carabids captured were Cychrus tuberculatus, Scaphinotus angusticollis and Omus dejeani, whereas the smallest carabid beetles captured were Notiophilus sylvaticus, Amara idahoensis, and Harpalus animosus.

Table 2. General data for sites sampled during 2005. Site Number of No. traps Total Average species sampled carabids seasonal over year catch 2nd growth sites

G1 13 229 2169 9.472 G2 14 185 692 3.741 G3L 13 155 1142 7.368 G3S 12 173 785 4.538 S1 9 59 598 10.136

Old growth sites G4 10 178 880 4.944 G5 12 123 898 7.301 L 10 116 699 6.026 M 12 187 973 5.203 O 9 74 523 7.068 Sm 9 135 418 3.096 C 8 58 123 2.121

Table 3. Total number of each carabid species caught June-September by location within group retention sites. Location G1 G2 G3L G3S G4 G5 L M Sm Out 10 11 9 8 7 8 7 8 8 5to10mOut 12 10 10 12 8 7 6 10 8

Edge 12 11 10 10 9 11 6 9 8 5to10mIn 12 11 9 9 In 12 11 13 12 9 8 9 10 8

19 Table 4. Total number of each carabid species caught May-September in each of the 11 sites. Species Name S1 G1 G2 G3L G3S G4 G5 L M Sm O C

Amara idahoensis 0 0 0 0 010 0 0 0 0 0 Cychrus tuberculatus 5 56 11 22 28 23 15 19 15 12 12 1 Harpalus animosus 0 0 1 0 000 0 2 0 0 0 Notiophilus sylvaticus 0 0 4 4 003 0 0 0 0 0 Omus dejeani 2 94 3 12 900 0 0 0 0 0 Promecognathus crassus 11 169 39 15 102 0 0 0 0 0 0 0 Pterostichus algidus 0 7 165 52 9214 0 0 0 0 0 Pterostichus amethystinus 0 19 7 11 904 1 5 2 3 1 Pterostichus castaneus 0 3 0 0 000 2 5 0 0 0 Pterostichus crenicollis 143 724 83 152 194 231 221 134 205 32 74 35 Pterostichus herculaneus 168 55 199 358 114 12 2 50 79 57 33 17 Pterostichus lama 12 47 32 19 26 21 72 21 21 49 33 56 Pterostichus pumilus pumilus 0 0 0 0 001 0 1 0 0 1 Scaphinotus angulatus 3 16 9 8 16 15 10 7 18 6 3 0 Scaphinotus angusticollis 252 876 124 439 269 523 421 437 550 232 336 9 Scaphinotus larvae 0 2 1 1 100 0 1 0 0 0 Scaphinotus marginatus 0 29 2 2 33345 17 19 21 12 0 Zacotus matthewsii 2 72 12 47 51990 11 52 7 17 3

20 Table 5. Presence/absence of the 17 carabid species in the 2nd growth sites. Carabid species are listed in order of decreasing overall abundance. Species Name S1 G1 G2 G3L G3S G4 G5 M L Sm O C N Scaphinotus angusticollis * * * * * * * * * * * * 4468 Pterostichus crenicollis * * * * * * * * * * * * 2228 Pterostichus herculaneus * * * * * * * * * * * * 1144 Pterostichus lama * * * * * * * * * * * * 409 Zacotus matthewsii * * * * * * * * * * * * 337 Promecognathus crassus * * * * * 336 Pterostichus algidus * * * * * * 249 Cychrus tuberculatus * * * * * * * * * * * * 219 Scaphinotus marginatus * * * * * * * * * * 183 Omus dejeani * * * * * 120 Scaphinotus angulatus * * * * * * * * * * * 111 Pterostichus amethystinus * * * * * * * * * * 62 Notiophilus sylvaticus * * * 11 Pterostichus castaneus * * * 10 Scaphinotus larvae * * * * * 6 * * * 3 Pterostichus pumilus pumilus Harpalus animosus * * 3 Amara idahoensis * 1

21

Table 6. Proportion of each carabid species by site. Most common species are outlined in bold.

Species Name S1 G1 G2 G3L G3S G4 G5 L M Sm O C Amara idahoensis 0.000 0.000 0.000 0.000 0.000 0.001 0.000 0.000 0.000 0.000 0.000 0.000 Cychrus tuberculatus 0.008 0.026 0.016 0.019 0.036 0.026 0.017 0.027 0.015 0.029 0.023 0.008 Harpalus animosus 0.000 0.000 0.001 0.000 0.000 0.000 0.000 0.000 0.002 0.000 0.000 0.000 Notiophilus sylvaticus 0.000 0.000 0.006 0.004 0.000 0.000 0.003 0.000 0.000 0.000 0.000 0.000 Omus dejeani 0.003 0.043 0.004 0.011 0.011 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Promecognathus 0.018 0.078 0.056 0.013 0.130 0.000 0.000 0.000 0.000 0.000 0.000 0.000 crassus Pterostichus algidus 0.000 0.003 0.238 0.046 0.011 0.002 0.016 0.000 0.000 0.000 0.000 0.000 Pterostichus 0.000 0.009 0.010 0.010 0.011 0.000 0.004 0.001 0.005 0.005 0.006 0.008 amethystinus Pterostichus castaneus 0.000 0.001 0.000 0.000 0.000 0.000 0.000 0.003 0.005 0.000 0.000 0.000 Pterostichus 0.239 0.334 0.120 0.133 0.247 0.263 0.246 0.192 0.211 0.077 0.141 0.285 crenicollis Pterostichus 0.281 0.025 0.288 0.313 0.145 0.014 0.002 0.072 0.081 0.136 0.063 0.138 herculaneus Pterostichus lama 0.020 0.022 0.046 0.017 0.033 0.024 0.080 0.030 0.022 0.117 0.063 0.455 Pterostichus pumilus 0.000 0.000 0.000 0.000 0.000 0.000 0.001 0.000 0.001 0.000 0.000 0.008 pumilus Scaphinotus angulatus 0.005 0.007 0.013 0.007 0.020 0.017 0.011 0.010 0.018 0.014 0.006 0.000 Scaphinotus 0.421 0.404 0.179 0.384 0.343 0.594 0.469 0.625 0.565 0.555 0.642 0.073 angusticollis Scaphinotus larvae 0.000 0.001 0.001 0.001 0.001 0.000 0.000 0.000 0.001 0.000 0.000 0.000 Scaphinotus 0.000 0.013 0.003 0.002 0.004 0.038 0.050 0.024 0.020 0.050 0.023 0.000 marginatus Zacotus matthewsii 0.003 0.033 0.017 0.041 0.006 0.022 0.100 0.016 0.053 0.017 0.033 0.024

22 Table 7. Proportion of each carabid species captured between May and September 2005 in 2nd growth sites G1 and G2, by location within sites. Species Name G1 G1 G1 G1 G1 G2 G2 G2 G2 5to10mI 5to10m Edge In Out 5to10m Edge In Out n Out Out Amara idahoensis 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Cychrus tuberculatus 0.019 0.053 0.006 0.021 0.037 0.005 0.034 0.008 0.029 Harpalus animosus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.010 Notiophilus sylvaticus 0.000 0.000 0.000 0.000 0.000 0.005 0.000 0.008 0.010 Omus dejeani 0.017 0.019 0.057 0.057 0.086 0.000 0.014 0.004 0.000 Promecognathus crassus 0.055 0.046 0.145 0.082 0.074 0.016 0.062 0.074 0.076 Pterostichus algidus 0.000 0.000 0.013 0.004 0.000 0.270 0.267 0.180 0.286 Pterostichus amethystinus 0.009 0.010 0.016 0.001 0.031 0.005 0.000 0.000 0.057 Pterostichus castaneus 0.000 0.002 0.006 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus crenicollis 0.247 0.476 0.249 0.342 0.344 0.200 0.089 0.078 0.124 Pterostichus herculaneus 0.036 0.012 0.035 0.021 0.031 0.265 0.281 0.293 0.324 Pterostichus lama 0.021 0.019 0.028 0.010 0.074 0.011 0.068 0.063 0.038 Pterostichus pumilus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 pumilus Scaphinotus angulatus 0.002 0.017 0.000 0.010 0.000 0.000 0.007 0.027 0.010 Scaphinotus angusticollis 0.514 0.320 0.401 0.407 0.294 0.211 0.144 0.234 0.038 Scaphinotus larvae 0.002 0.000 0.003 0.000 0.000 0.005 0.000 0.000 0.000 Scaphinotus marginatus 0.013 0.014 0.013 0.014 0.012 0.005 0.007 0.000 0.000 Zacotus matthewsii 0.064 0.012 0.028 0.031 0.018 0.000 0.027 0.031 0.000

23

Table 8. Proportion of each carabid species captured between May and September 2005 in 2nd growth sites G3, by location within both large and small patches. Species Name G3 G G3 G3 G3 G3 G3 G3 G3 Large Large Large Large Large Small Small Small Small 5to10m 5to10m Edge In Out 5to10m Edge In Out In Out Out Amara idahoensis 0 0 0 0 0 0 0 00 Cychrus tuberculatus 0.010 0.073 0.011 0.013 0.042 0.020 0.043 0.042 0.032 Harpalus animosus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Notiophilus sylvaticus 0.000 0.000 0.000 0.007 0.000 0.000 0.000 0.000 0.000 Omus dejeani 0.013 0.009 0.043 0.004 0.014 0.005 0.011 0.012 0.032 Promecognathus crassus 0.006 0.036 0.043 0.005 0.028 0.113 0.141 0.090 0.365 Pterostichus algidus 0.044 0.082 0.250 0.005 0.042 0.015 0.016 0.009 0.000 Pterostichus amethystinus 0.003 0.009 0.022 0.005 0.056 0.015 0.016 0.009 0.000 Pterostichus castaneus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus crenicollis 0.060 0.209 0.076 0.154 0.250 0.488 0.200 0.144 0.159 Pterostichus herculaneus 0.381 0.345 0.304 0.266 0.347 0.089 0.114 0.204 0.111 Pterostichus lama 0.016 0.073 0.033 0.005 0.000 0.015 0.032 0.030 0.111 Pterostichus pumilus pumilus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Scaphinotus angulatus 0.010 0.018 0.000 0.004 0.014 0.005 0.022 0.030 0.016 Scaphinotus angusticollis 0.378 0.145 0.207 0.488 0.208 0.227 0.405 0.410 0.175 Scaphinotus larvae 0.000 0.000 0.000 0.002 0.000 0.000 0.000 0.003 0.000 Scaphinotus marginatus 0.000 0.000 0.000 0.004 0.000 0.005 0.000 0.006 0.000 Zacotus matthewsii 0.079 0.000 0.011 0.038 0.000 0.005 0.000 0.012 0.000

24 Table 9. Proportion of each carabid species captured between May and September 2005 in old growth sites G4&G5, by location. Species Name G4 G4 G4 G4 G G5 G5 G5 G5 5to10m Edge In Out 5to10m 5to10m Edge In Out Out In Out Amara idahoensis 0.000 0.000 0.000 0.016 0.000 0.000 0.000 0.000 0.000 Cychrus tuberculatus 0.044 0.029 0.015 0.047 0.006 0.029 0.024 0.011 0.042 Harpalus animosus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Notiophilus sylvaticus 0.000 0.000 0.000 0.000 0.006 0.010 0.005 0.000 0.000 Omus dejeani 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Promecognathus crassus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus algidus 0.000 0.003 0.003 0.000 0.081 0.000 0.005 0.000 0.000 Pterostichus amethystinus 0.000 0.000 0.000 0.000 0.000 0.029 0.005 0.000 0.000 Pterostichus castaneus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus crenicollis 0.372 0.186 0.278 0.344 0.255 0.275 0.193 0.245 0.396 Pterostichus herculaneus 0.027 0.007 0.013 0.031 0.000 0.000 0.000 0.003 0.021 Pterostichus lama 0.018 0.010 0.005 0.219 0.050 0.186 0.075 0.069 0.063 Pterostichus pumilus pumilus 0.000 0.000 0.000 0.000 0.000 0.000 0.005 0.000 0.000 Scaphinotus angulatus 0.053 0.007 0.015 0.016 0.006 0.000 0.005 0.019 0.021 Scaphinotus angusticollis 0.442 0.694 0.604 0.328 0.416 0.373 0.547 0.499 0.271 Scaphinotus larvae 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Scaphinotus marginatus 0.027 0.039 0.045 0.000 0.019 0.098 0.024 0.051 0.167 Zacotus matthewsii 0.018 0.026 0.023 0.000 0.161 0.000 0.113 0.104 0.021

25 Table 10. Proportion of each carabid species captured between May and September 2005 in old growth VRAM treatments Sm & M, by location. Species Name M M MIn MOut Sm Sm SmIn SmOut 5to10m Edge 5to10m Edge Out Out Amara idahoensis 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Cychrus tuberculatus 0.023 0.020 0.010 0.021 0.056 0.027 0.020 0.037 Harpalus animosus 0.000 0.000 0.000 0.021 0.000 0.000 0.000 0.000 Notiophilus sylvaticus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Omus dejeani 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Promecognathus crassus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus algidus 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 Pterostichus amethystinus 0.016 0.000 0.004 0.011 0.037 0.000 0.000 0.000 Pterostichus castaneus 0.023 0.004 0.002 0.000 0.000 0.000 0.000 0.000 Pterostichus crenicollis 0.388 0.084 0.224 0.232 0.111 0.045 0.060 0.167 Pterostichus herculaneus 0.078 0.068 0.038 0.347 0.315 0.225 0.015 0.222 Pterostichus lama 0.039 0.020 0.010 0.063 0.093 0.171 0.045 0.296 Pterostichus pumilus pumilus 0.000 0.004 0.000 0.000 0.000 0.000 0.000 0.000 Scaphinotus angulatus 0.031 0.000 0.028 0.000 0.019 0.018 0.005 0.037 Scaphinotus angusticollis 0.364 0.715 0.600 0.263 0.333 0.468 0.759 0.204 Scaphinotus larvae 0.000 0.004 0.000 0.000 0.000 0.000 0.000 0.000 Scaphinotus marginatus 0.008 0.020 0.026 0.000 0.037 0.027 0.075 0.019 Zacotus matthewsii 0.031 0.060 0.058 0.042 0.000 0.018 0.020 0.019

26

Table 11. Proportion of each carabid species captured between June and September 2005 in old growth sites VRAM treatment L, by location. Species Name L5to10m L5to10m L L L In Out Edge In Out Amara idahoensis 0.000 0.000 0.000 0.000 0.000 Cychrus tuberculatus 0.004 0.045 0.094 0.020 0.103 Harpalus animosus 0.000 0.000 0.000 0.000 0.000 Notiophilus sylvaticus 0.000 0.000 0.000 0.000 0.000 Omus dejeani 0.000 0.000 0.000 0.000 0.000 Promecognathus crassus 0.000 0.000 0.000 0.000 0.000 Pterostichus algidus 0.000 0.000 0.000 0.000 0.000 Pterostichus amethystinus 0.004 0.000 0.000 0.000 0.000 Pterostichus castaneus 0.000 0.000 0.000 0.007 0.000 Pterostichus crenicollis 0.102 0.358 0.156 0.241 0.138 Pterostichus herculaneus 0.024 0.134 0.344 0.027 0.172 Pterostichus lama 0.020 0.075 0.063 0.007 0.172 Pterostichus pumilus pumilus 0.000 0.000 0.000 0.000 0.000 Scaphinotus angulatus 0.004 0.015 0.000 0.017 0.000 Scaphinotus angusticollis 0.812 0.373 0.328 0.619 0.345 Scaphinotus larvae 0.000 0.000 0.000 0.000 0.000 Scaphinotus marginatus 0.004 0.000 0.016 0.048 0.034 Zacotus matthewsii 0.024 0.000 0.000 0.014 0.034

27 Table 12. Average catches of each carabid species for May-September For Size of Patch (C=control, L= large, M =medium, Sm=small) 2nd growth sites Old growth sites Site Name S1 G1 G3L G3Sm G2 O1 G5 L M G4 Sm C1 Size of Patch C L L Sm Sm C L L M Sm Sm C Amara idahoensis 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.01 0.00 0.00 Cychrus 0.08 0.23 0.14 0.14 0.06 0.17 0.11 0.15 0.08 0.12 0.09 0.02 tuberculatus Harpalus 0.00 0.00 0.00 0.00 0.01 0.00 0.00 0.00 0.02 0.00 0.00 0.00 animosus Notiophilus 0.00 0.00 0.02 0.00 0.02 0.00 0.02 0.00 0.00 0.00 0.00 0.00 sylvaticus Omus dejeani 0.03 0.37 0.08 0.05 0.02 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Promecognathus 0.18 0.69 0.10 0.60 0.21 0.00 0.00 0.00 0.00 0.00 0.00 0.00 crassus Pterostichus 0.00 0.03 0.36 0.05 0.88 0.00 0.14 0.00 0.00 0.01 0.00 0.00 algidus Pterostichus 0.00 0.09 0.07 0.05 0.04 0.04 0.04 0.01 0.02 0.00 0.02 0.02 amethystinus Pterostichus 0.00 0.01 0.00 0.00 0.00 0.00 0.00 0.01 0.03 0.00 0.00 0.00 castaneus Pterostichus 2.42 3.05 0.90 1.07 0.45 0.99 1.72 1.06 0.98 1.09 0.23 0.61 crenicollis Pterostichus 2.89 0.24 2.20 0.56 1.05 0.45 0.02 0.44 0.41 0.06 0.40 0.29 herculaneus Pterostichus lama 0.21 0.21 0.12 0.16 0.16 0.45 0.55 0.20 0.11 0.11 0.35 0.97 Pterostichus 0.00 0.00 0.00 0.00 0.00 0.00 0.01 0.00 0.01 0.00 0.00 0.02

28 pumilus pumilus Scaphinotus 0.05 0.07 0.05 0.08 0.04 0.04 0.09 0.06 0.08 0.08 0.04 0.00 angulatus Scaphinotus 4.24 3.72 2.58 1.33 0.62 4.51 3.31 3.68 2.55 2.55 1.64 0.16 angusticollis Scaphinotus 0.00 0.12 0.01 0.01 0.01 0.16 0.37 0.13 0.09 0.15 0.15 0.00 marginatus Zacotus matthewsii 0.03 0.31 0.28 0.02 0.06 0.23 0.68 0.09 0.26 0.09 0.04 0.05

All carabids 10.14 9.14 6.92 4.10 3.64 7.04 7.07 5.84 4.63 4.26 2.96 2.13 average catch No. Species 9 13 13 12 14 9 12 10 12 10 9 8

29

Patterns of abundance

Seasonal patterns of abundance showed general peaks of abundance in May and September in most sites, for both total carabids pooled and for Scaphinotus angusticollis, the most commonly captured beetle (Figures 3-8). These figures show the catches of total carabids and S. angusticollis per patch trap, rather than averaged for the entire site. In both cases, there was a clear pattern of higher catches in the large patch sites over the smaller patch sites, for both old growth and 2nd growth sites. For 2nd growth sites, abundance was higher in the large patch sites G1 and G3Large, followed by the 2nd growth control, with lowest abundance overall in the small patch sites, G3 small and G2 (Figures 3&6). For old growth sites, catches were highest in the large patch site G5, followed by the old growth control, then the small patch site G4, and lowest overall in the clearcut (Figures 4&7). For the VRAM treatments, abundance varied from highest in the large patch treatment (L), next in the medium (M) treatment, then the old growth control, followed by the small patch treatment (Sm) and lowest overall in the clearcut (Figures 5&8). The next set of graphs show the changes in abundance of the most commonly captured species along the transects moving from the centre of patches, across the edge, out to the matrix (Figures 9-21). For some species, catches appeared to increase at the edge of patches, or at locations 5-10 m out from the edge into the matrix. This was the case for Cychrus tuberculatus in both 2nd and old growth sites (Figures 9&10), P. crassus in site G1 (Figure 11), P. crenicollis in 2nd growth sites (Figure 13), and P. herculaneus in old growth sites (Figure 16). Abundance of Zacotus matthewsii showed peaks 5-10 m from the edge inside of patches in the large patch sites in 2nd growth (Figure 19), and this was also apparent in the large patches of site G3 for P. herculaneus (Figure 15). The forest specialist species (S. angusticollis, P. crenicollis and Z. matthewsii) showed an expected increase inside of patches as compared to in the matrices, while generalist species such as P. lama, P. crassus, C. tuberculatus and P. herculaneus generally showed no clear differences in abundance among patch and matrix locations. The forest specialist species were generally more abundant in the large patches over small patches in both old growth and 2nd growth sites. P. crenicollis was not well retained in the small

30 patch treatment of the VRAM site, as compared to the larger patch size treatments in the VRAM, and was also not retained well in the 2nd growth small patches in site G3 and G2, as compared with sites G1 and large patches of site G3 (Figures 12&13). Zacotus was similarly better retained in the large patch site G5, than in the other old growth sites, and was also far less well retained in the 2nd growth small patches in sites G2 and G3 than in the large patches of sites G3 and G1 (Figures 18&19). S. angusticollis was well retained in the patches all of the old growth sites, but in 2nd growth site, the same pattern of poor retention in the small patch 2nd growth sites, was apparent (Figures 20&21). Although retention of forest specialists was generally better in old growth patches than 2nd growth patches, retention was always lowest overall in the small patch VRAM sites than in the old growth sites with larger patches.

31 Figure 3. Average catches of all carabid beetle species pooled by date in 2nd growth sites. Catches are given per patch trap for the group retention sites.

2nd growth carabid catches by Patch Size 1.4 G1 1.2 G2

p G3 Large G3 Small 1.0 S

0.8

0.6

0.4 Average no. beetles per tra per beetles no. Average 0.2

0.0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

Figure 4. Average catches of all carabid beetle species pooled by date in old growth sites G4 and G5. Catches in VRAM clearcut and old growth treatments are shown for comparison. Catches are given per patch trap for the group retention sites.

Old Growth carabid Catches by Patch Size

1.2

G4 1.0 G5 p O C 0.8

0.6

0.4

Average no. beetles per tra per beetles no. Average 0.2

0.0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

32 Figure 5. Average catches of all carabid beetle species pooled by date in old growth VRAM treatments. Catches are given per patch trap for the group retention sites.

VRAM carabid Catches by Patch Size 1.2

SM M 1.0 p L O C 0.8

0.6

0.4

Average no. beetles per tra per beetles no. Average 0.2

0.0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

Figure 6. Average catches of Scaphinotus angusticollis by date in 2nd growth sites. Catches are given per patch trap for the group retention sites.

2nd growth S. angusticollis catches by Patch Size 16 G1 14 G2

p G3 Large 12 G3 Small S 10

8

6

4 Average no. beetles per tra per beetles no. Average 2

0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

33 Figure 7. Average catches of Scaphinotus angusticollis by date in old growth sites G4 and G5. Catches in VRAM clearcut and old growth treatments are shown for comparison. Catches are given per patch trap for the group retention sites.

Old Growth S. angusticollis Catches by Patch Size 16

14 G4 G5 p O 12 C

10

8

6

4 Average no. beetles per tra per beetles no. Average 2

0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

Figure 8. Average catches of Scaphinotus angusticollis by date in old growth VRAM treatments. Catches are given per patch trap for the group retention sites.

VRAM S. angusticollis Catches by Patch Size 16

SM 14 M

p L 12 O C 10

8

6

4 Average no. beetles per tra per beetles no. Average 2

0 May-05 Jun-05 Jul-05 Aug-05 Sep-05

34 Figure 9. Average yearly catches of Cychrus tuberculatus by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites C. tuberculatus

0.30 Into cut matrices G5 Into forest patches L 0.25 G4 Sm M 0.20

0.15

0.10 Mean no. beetles per trap per beetles no. Mean 0.05

0.00 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 10. Average yearly catches of Cychrus tuberculatus by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites C. tuberculatus

1.40 Into cut matrices Into forest patches 1.20

G1 1.00 G3L G2 0.80 G3Sm

0.60

0.40 Mean no. beetles per trap per beetles no. Mean 0.20

0.00 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

35 Figure 11. Average yearly catches of Promognathus crassus by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites P. crassus

1.40 Into cut matrices Into forest patches 1.20

1.00

0.80

G1 0.60 G3L G2 0.40 G3Sm Mean no. beetles per trap per beetles no. Mean 0.20

0.00 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Figure 12. Average yearly catches of Pterostichus crenicollis by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites P. crenicollis

3.0 G5 Into cut matrices Into forest patches L 2.5 G4 Sm M 2.0

1.5

1.0 Mean no. beetles per trap per beetles no. Mean 0.5

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

36 Figure 13. Average yearly catches of Pterostichus crenicollis by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites P. crenicollis

6 G1 Into cut matrices Into forest patches G3L 5 G2 G3Sm 4

3

2 Mean no. beetles per trap per beetles no. Mean 1

0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Figure 14. Average yearly catches of Pterostichus lama by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites P. lama

1.00 Into cut matrices Into forest patches 0.90

0.80

0.70

0.60 G5 L 0.50 G4 0.40 Sm M 0.30

Mean no. beetles per trap per beetles no. Mean 0.20

0.10

0.00 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

37 Figure 15. Average yearly catches of Pterostichus lama by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites P. lama

0.30 Into cut matrices Into forest patches G1 0.25 G3L G2 G3Sm 0.20

0.15

0.10 Mean no. beetles per trap per beetles no. Mean 0.05

0.00 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Figure 16. Average yearly catches of Pterostichus herculaneus by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites P.herculaneus

1.2 Into cut matrices Into forest patches G5 L 1.0 G4 Sm 0.8 M

0.6

0.4 Mean no. beetles per trap per beetles no. Mean 0.2

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

38 Figure 17. Average yearly catches of Pterostichus herculaneus by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites P. herculaneus

4.50 Into cut matrices Into forest patches 4.00

3.50

3.00

2.50 G1 G3L 2.00 G2 G3Sm 1.50

1.00 Mean no. beetles per trap per beetles no. Mean

0.50

0.00 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Figure 18. Average yearly catches of Zacotus matthewsii by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Z. matthewsii

1.4 Into forest patches G5 Into cut matrices L 1.2 G4 Sm 1.0 M

0.8

0.6

0.4 Mean no. beetles per trap per beetles no. Mean 0.2

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

39 Figure 19. Average yearly catches of Zacotus matthewsii by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Z. matthewsii

1.0 Into cut matrices Into forest patches G1 0.9 G3L G2 0.8 G3Sm 0.7

0.6

0.5

0.4

0.3

Mean no. beetles per trap per beetles no. Mean 0.2

0.1

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Figure 20. Average yearly catches of Scaphinotus angusticollis by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old growth S. angusticollis

9 G5 Into cut matrix Into forest patches L 8 G4 Sm 7 M

6

5

4

3

Mean no. beetles per trap per beetles no. Mean 2

1

0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

40 Figure 21. Average yearly catches of Scaphinotus angusticollis by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites S. angusticollis

8 G1 Into cut matrices Into forest patches G3L 7 G2 G3Sm 6

5

4

3

2 Mean no. beetles per trap per beetles no. Mean

1

0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

Statistical analysis

Individual species differences among treatments and locations within group retention types.

Paired t-tests to compare catches among patches and matrices within group VR sites. I focussed these comparisons on forest specialist species, since these are the species we are most interested in retaining within patches. Results are listed in Table 13 below. In all cases, forest specialists were more abundant in the patches than in the matrices surrounding patches, but this could not always be discerned statistically (using P<0.05) when we examined the overall mean catches per patch trap versus mean catches per matrix trap for the 4 or 5 months of trapping. 2nd growth: For Scaphinotus angusticollis, catches were significantly greater in the patches of all the sites than in the matrices, except for the large patches of site G3. For P. crenicollis, catches were significantly greater in the patches of sites G1 and G3S than the

41 surrounding matrices, but did not differ significantly between the patches and matrices of sites G2 and G3L. For Zacotus matthewsii, catches were significantly greater in the patches of sites G1 and G3L than the surrounding matrices, but did not differ significantly between the patches and matrices of sites G2 and G3S.

Old growth: For Scaphinotus angusticollis, catches did not differ significantly among the patches and matrices of any of the sites, except for site Sm, where they were significantly greater in the patches than the matrix. For P. crenicollis, catches were significantly greater in the patches of sites G4, G5, and M than in the surrounding matrices, but did not differ significantly between the patches and matrices of sites L and Sm. For Zacotus matthewsii, catches were significantly greater in the patches of sites G4, G5 and M, than in the matrices, but did not differ significantly among the patches and matrices of the VRAM treatments Sm and L.

Table 13. Results of paired t-tests to compare mean abundance of three forest specialists in patches versus matrices of group retention sites, by patch size, and forest type (2nd growth or old growth). Mean abundance for each species by site and location for the whole season are given, together with values of T, df, and P. Significant results (P<0.05) are shown in bold.

a. Scaphinotus angusticollis Scaphinotus angusticollis 2nd Growth Sites Patch mean Matrix mean t df P G1 6.001 1.069 4.752 4 0.009 G2 1.035 0.104 3.606 4 0.023 G3L 6.75 0.6 2.658 4 0.056 G3S 2.283 0.44 2.773 4 0.05

Old Growth Sites Patch mean Matrix mean t df P G4 4.498 0.7 2.179 4 0.095 G5 6.157 0.85 2.222 4 0.09 L 7.545 0.5 2.81 3 0.067 M 5.095 0.556 2.669 4 0.056 Sm 4.152 0.313 3.599 3 0.037

42 b. Pterostichus crenicollis Pterostichus crenicollis 2nd Growth Sites Patch mean Matrix meant df P G1 5.012 1.264 6.528 4 0.003 G2 0.358 0.32 6.887 4 0.002 G3L 2.125 0.687 2.294 4 0.083 G3S 0.8 0.393 3.992 4 0.016

Old Growth Sites Patch mean Matrix meant df P G4 1.952 0.592 4.022 4 0.016 G5 2.682 0.98 5.097 4 0.007 L 2.538 0.2 2.686 3 0.075 M 1.977 0.532 4.253 4 0.013 Sm 0.309 0.266 0.662 3 0.555 c. Zacotus matthewsii Zacotus matthewsii 2nd Growth Sites Patch mean Matrix meant df P G1 0.468 0.069 2.98 4 0.041 G2 0.138 0 2.431 4 0.072 G3L 0.525 0 3.628 4 0.022 G3S 0.067 0 2.138 4 0.099

Old Growth Sites Patch mean Matrix meant df P G4 0.165 0 2.915 4 0.043 G5 1.2 0.05 3.104 4 0.036 L 0.159 0.05 2.92 3 0.062 M 0.646 0.106 2.814 4 0.048 Sm 0.083 0.028 0.577 3 0.604

T-tests to compare catches among matrices of old growth VRAM group treatments and clearcut catches. Again, these comparisons were focussed on forest specialist species. Results are listed in Table 14 below. T-tests showed that abundances of S. angusticolli and P. crenicollis did not differ significantly in each of the Small, Medium or Large site

43 matrices when these catches were compared with the seasonal clearcut catches of this species. In the case of Z. matthewsii, catches between the matrices of both the Small and Medium treatments did not differ significantly to the catches of this species in the clearcut treatment, but catches of Zacotus were significantly greater in the matrices of the Medium Group VR sites than in the clearcut.

Table 14. Results of t-tests to compare catches of forest specialists among matrices of old growth VRAM group treatments and clearcut catches. Mean abundance for each species within matrices of the three VRAM group retention treatments and within the clearcut for the whole season are given, together with values of T, df, and P. Significant results (P<0.05) are shown in bold.

Scaphinotus angusticollis

Matrix Clearcut t df P L 0.5 0.158 1.068 3 0.364

M 0.634 0.158 1.118 3 0.345

Sm 0.313 0.158 1.319 3 0.279

Pterostichus crenicollis

Matrix Clearcut t df P L 0.2 0.608 -1.275 3 0.292

M 0.498 0.608 -0.447 3 0.685

Sm 0.266 0.608 -1.228 3 0.307

Zacotus matthewsii

Matrix Clearcut t df P L 0.05 0.051 -0.018 3 0.987

M 0.133 0.051 5.456 3 0.012

Sm 0.028 0.051 -0.522 3 0.638

44

Paired t-tests to compare catches among differently sized patches. Results are listed in Table 15 below. Note that catches of forest specialists were always greater in the large patches over the small patches (except for Zacotus matthewsii which was more abundance in the M VRAM treatment than the L VRAM treatment) in all of the following comparisons, but using a P<0.05 as our significance level, it was not always possible to find a statistically significant difference. a) Catches within large patches and small patches in site G3: abundance of both S. angusticollis and P. crenicollis did not differ significantly among the differently- sized patches, but Z. matthewsii was less abundant in the smaller patches of site G3 than in the larger patches in the same site. b) Catches from small patches of site G2 with large patches of site G1: abundances of S. angusticollis , P. crenicollis and Z. matthewsii were each significantly greater in the large patches of site G1 than in the smaller patches of site G2. c) Catches in small patches of site G4 with large patches of site G5: abundance of S. angusticollis and P. crenicollis did not differ significantly among small patches in site G4 and large patches in site G5, but Z. matthewsii was significantly more abundant in the large patches of G5 than in small patches of G4. d) Catches among small, medium and large patches of the VRAM site: abundance of S. angusticollis and Z. matthewsii did not differ significantly between Large patches and Small patches, between Small and Medium patches, or between Large and Medium sized patches. Catches of P. crenicollis did not differ significantly among Large and Medium size patches, but were significantly less abundant in Small patches than in both Large and Medium patches.

Table 15. Results of paired t-tests to compare catches among differently sized patches (large vs. small for 2nd growth sites, large vs medium vs. small for old growth sites. Mean abundance for each species within the patches for the whole season are given, together with values of T, df, and P. Significant results (P<0.05) are shown in bold. a. Scaphinotus angusticollis Scaphinotus angusticollis 2nd Growth Sites

45 Small patch Large patch t df P G2 G1 1.035 6.001 4.031 4 0.016 G3S G3L 2.283 6.75 2.549 4 0.063

Old Growth Sites Small patch Large patch t df P G4 G5 4.498 6.157 2.118 4 0.102 M L 5.095 7.545 3>0.1 Sm L 4.152 7.545 1.067 3 0.364 M Sm 5.095 4.152 3>0.1 b. Pterostichus crenicollis Pterostichus crenicollis 2nd Growth Sites Small patch Large patch t df P G2 G1 0.358 5.012 6.887 4 0.002 G3S G3L 0.8 2.125 1.941 4 0.124

Old Growth Sites Small patch Large patch t df P G4 G5 1.952 2.682 1.35 4 0.248 M L 1.977 2.538 0.9 3 0.434 Sm L 0.309 2.538 3.235 3 0.048 Sm M 0.309 1.977 4.615 3 0.019 c. Zacotus matthewsii Zacotus matthewsii 2nd Growth Sites Small patch Large patch t df P G2 G1

46 0.138 0.468 3.885 4 0.018 G3S G3L 0.067 0.525 2.972 4 0.041

Old Growth Sites Small patch Large patch t df P G4 G5 0.165 1.2 3.088 4 0.037 M L 0.646 0.159 2.37 3 0.062 Sm L 0.083 0.159 0.838 3 0.464 Sm M 0.083 0.646 2.477 3 0.092

Analysis of differences among group VR patch catches and control site catches of forest specialists. We used one-way ANOVA and Tukey’s post-hoc test to compare the mean monthly catches of the two forest specialists, S.angusticollis and P. crenicollis, in the patch locations (“In” traps) within each of the group retention sites, with the catches in the relevant control. The control site was S1 for comparison with the 2nd growth group VR sites, and was O, the old growth VRAM site, for comparison with the old growth group VR sites. Comparison of catches of Zacotus matthewsii were made among sites for all dates pooled since the monthly catches of this species were generally lower than for the other species.

2nd growth sites: Catches of S. angusticollis varied significantly among sites for all months, May to September, with the exception of August, when no significant differences could be found (P>0.05). In May there were higher catches in the large patches in sites G1 and G3 than in the small patches of sites G2 and G3, but patch catches did not differ from the 2nd growth control catches (F=7.656, df=4,51, P=0.000). The same findings were true of catches of S. angusticollis in June (F=2.955, df=4,50, P=0.029). In July, highest catches overall were from the large patches of site G1, which also had higher catches than in the 2nd growth control site (F=2.974, df=4,49, P=0.028). In September, catches from the small patches in site G2 were lower than from the large

47 patches in sites G1 and G3, but catches did not differ significantly from the 2nd growth control in any of the sites (F=3.88, df=4,49, P=0.007). Catches of P.crenicollis varied significantly among sites for all months except May and August (P>0.05). In each of June, July, and September, catches were higher in the large patches of site G1 than in the small patches of sites G2 and G3 (June: F=3.391, df=4,50, P=0.016; July: F=4.389, df=4,49, P=0.004; September: F=6.32, df=4,48, P=0.000). Again, post-hoc testing did not show that catches of this species differed significantly among the patches of the group VR sites and the 2nd growth control on any date. Catches of Zacotus varied significantly among sites, with highest catches in the large patches of site G3 and G1 overall: catches in these patches was also significantly higher than in the 2nd growth control (F=6.058, df=4,20, P=0.002).

Old growth sites: Catches of S. angusticollis did not vary among sites for any months (P>0.05) except for August, when there were significantly greater numbers caught in the large patches of the VRAM treatment L, than in the patches of all other sites as well as in the old growth control (F=5.812, df=5,57, P=0.0000). Catches of P. crenicollis did not vary among the patches of the old growth group VR sites and the old-growth control during any of the sampling months, May to September (P>0.05 for all months). Catches of Zacotus varied significantly among sites, with highest catches in the large patches of site G5 overall: catches in these patches was also significantly higher than in the old growth control (F=5.968, df=5,22, P=0.001).

General Linear Model to examine responses of individual species to locations within patch retention cuts. For the most commonly captured species, I used a General Linear model to examine the responses by species to different trap locations within group retentions sites, with the trap locations (In, Out, Edge, 5 to 10m in from Edge, 5 to 10m out from Edge) nested within sites. Results for 2nd growth and old growth sites are given in Tables 16& 17 below.

48 2nd growth sites: • For Zacotus matthewsi, sites (G1, G2, G3small and G3large) did not differ significantly, but locations nested within sites (In, Out, Edge, 5 to 10m in from Edge & 5 to 10m out from Edge) did differ significantly, , with highest numbers overall in the 5 to10m in from edge locations . • For Scaphinotus angusticollis, sites (G1, G2, G3small and G3large) did not differ significantly, but locations nested within sites (In, Out, Edge, 5 to 10m in from Edge & 5 to 10m out from Edge) did differ significantly, with higher abundance in the In and 5 to 10m out from edge locations. • For Pterostichus crenicollis, both (G1, G2, G3small and G3large) and locations within sites (In, Out, Edge, 5 to 10m in from Edge & 5 to 10m out from Edge) did differ significantly, with generally most caught in site G1, and most from inside patches, and in some sites, at the 5-10m from the edge of traps into the matrix locations. • For P. herculaneus, both sites (G1, G2, G3small and G3large) and locations within sites (In, Out, Edge, 5 to 10m in from Edge & 5 to 10m out from Edge) differed significantly, with highest numbers overall in the large patch traps of Site G3. • Finally, for P. lama, there were no significant differences in abundance for both sites (G1, G2, G3small and G3large) and locations within sites (In, Out, Edge, 5 to 10m in from Edge & 5 to 10m out from Edge).

Old growth sites: • For Zacotus matthewsi, sites (G4, G5, Sm, M, L) and locations within sites (In, Out, Edge, 10m in from Edge & 5 to 10m out from Edge) differed significantly, with highest numbers overall in Site G5, and higher numbers from In and 5 to 10m in from Edge traps than 5 to 10m out from Edge and Out traps. • For Scaphinotus angusticollis, sites (G4, G5, Sm, M, L) did not differ significantly, but locations nested within sites (In, Out, Edge, 5 to 10m in

49 from Edge & 5 to 10m out from Edge) did differ significantly, with higher abundance in the In and 5 to 10m in from Edge locations. • For Pterostichus crenicollis, catches differed significantly among sites (G4, G5, Sm, M, L) but not locations within sites (In, Out, Edge, 10m in from Edge & 5 to 10m out from Edge). Most P. crenicollis were caught in G5 overall. • For P. herculaneus, catches differed significantly among sites (G4, G5, Sm, M, L) but not locations within sites (In, Out, Edge, 10m in from Edge & 5 to 10m out from Edge). Sites G4 and G5 had lower abundance than the other sites. • Finally, for P. lama, catches differed significantly among sites (G4, G5, Sm, M, L) but not locations within sites (In, Out, Edge, 10m in from Edge & 5 to 10m out from Edge). Highest numbers of P. lama were caught in site G5 overall.

Table 16. Results of a general linear model to examine the responses of individual species to different trap locations within 2nd growth group retention sites. Trap locations (In, Out, Edge, 5 to 10m out from Edge, and 5 to 10m in from Edge) were nested within sites. In each case, the error term df=72. Species F df P Zacotus Site 2.138 4 0.085 matthewsii Location 0.603 3 0.615 Location(Site) 2.044 10 0.041 Scaphinotus Site 0.282 3 0.838 angusticollis Location 1.924 3 0.133 Location(Site) 2.119 11 0.029 Pterostichus Site 0.126 3 0.944 crenicollis Location 24.116 3 0.000 Location(Site) 4.587 11 0.000 Pterostichus Site 0.296 3 0.828 herculaneus Location 45.844 2 0.000 Location(Site) 4.419 12 0.000 Pterostichus Site 3 >0.05 lama

50 Location 3 >0.05 Location(Site) 11 >0.05

Table 17. Results of a general linear model to examine the responses of individual species to different trap locations within old growth group retention sites. Trap locations (In, Out, Edge, 5 to 10m out from Edge, and 5 to 10m in from Edge) were nested within sites. In each case, the error term df=79. Species F df P Zacotus Site 13.986 4 0.000 matthewsii Location 0.537 3 0.658 Location(Site) 2.942 14 0.001 Scaphinotus Site 0.573 3 0.67 angusticollis Location 2.601 4 0.042 Location(Site) 1.187 14 0.302 Pterostichus Site 7.15 4 0.000 crenicollis Location 1.839 3 0.147 Location(Site) 1.251 14 0.257 Pterostichus Site 5.163 4 0.001 herculaneus Location 3 0.997 Location(Site) 14 0.366 Pterostichus Site 6.686 4 0.000 lama Location 0.622 3 0.603 Location(Site) 2.124 14 0.019

51 Species-patch area curves

Using estimates of patch area, calculated from setting maps for all the sites, species-patch area curves were produced for Scaphinotus angusticollis, S. marginatus, Pterostichus crenicollis, P. lama, P. herculaneus and Zacotus matthewsii for both old growth sites and second growth sites (Figures 22-34). The number of each species caught per trap inside the patches of each group retention site, over the whole trapping season, was plotted against the relevant patch area. From these figures, it is apparent that for the forest specialists, S. angusticollis, P. crenicollis, and Zacotus matthewsii, there is clear response of better retention of these species with increasing patch size. The same response is also seen in the old growth sites for S. marginatus, another species found associated with forest. The two generalist species, P. lama and P.herculaneus were caught in low abundance in old-growth sites, but showed unclear patterns, or no change in abundance with increasing patch sizes.

52 Figure 23. Mean seasonal catch (beetles per trap) of Scaphinotus angusticollis (with associated standard error bars) plotted against patch area for old growth sites.

Scaphinotus angusticollis old growth

9

8

h 7

6

5

4

3

Average seasonal catc seasonal Average 2

1

0 0 10000 20000 30000 40000 50000 60000 Patch area (sq. meters)

Figure 24. Mean seasonal catch (beetles per trap) of Scaphinotus angusticollis (with associated standard error bars) plotted against patch area for second growth sites.

Scaphinotus angusticollis 2nd growth

9

8

h 7

6

5

4

3

Average seasonal catc seasonal Average 2

1

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

53 Figure 25. Mean seasonal catch (beetles per trap) of Scaphinotus marginatus (with associated standard error bars) plotted against patch area for old growth sites.

Scaphinotus marginatus old growth

0.7

0.6 h 0.5

0.4

0.3

0.2 Average seasonal catc seasonal Average

0.1

0 0 10000 20000 30000 40000 50000 60000 Patch area (sq. meters)

Figure 26. Mean seasonal catch (beetles per trap) of Scaphinotus marginatus (with associated standard error bars) plotted against patch area for second growth sites.

Scaphinotus marginatus 2nd growth 0.3

0.25 h

0.2

0.15

0.1 Average seasonal catc seasonal Average

0.05

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

54 Figure 27. Mean seasonal catch (beetles per trap) of Pterostichus crenicollis (with associated standard error bars) plotted against patch area for old growth sites.

Pterostichus crenicollis old growth

3.5

3 h 2.5

2

1.5

1 Average seasonal catc seasonal Average

0.5

0 0 10000 20000 30000 40000 50000 60000 Patch area (sq. meters)

Figure 28. Mean seasonal catch (beetles per trap) of Pterostichus crenicollis (with associated standard error bars) plotted against patch area for second growth sites.

Pterostichus crenicollis 2nd growth 7

6 h 5

4

3

2 Average seasonal catc seasonal Average

1

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

55 Figure 29. Mean seasonal catch (beetles per trap) of Pterostichus lama (with associated standard error bars) plotted against patch area for old growth sites.

Pterostichus lama old growth

1.00E+00

9.00E-01

8.00E-01 h 7.00E-01

6.00E-01

5.00E-01

4.00E-01

3.00E-01

2.00E-01 Average seasonal catc seasonal Average 1.00E-01

0.00E+00 0 10000 20000 30000 40000 50000 60000 -1.00E-01 Patch area (sq. meters)

Figure 30. Mean seasonal catch (beetles per trap) of Pterostichus lama (with associated standard error bars) plotted against patch area for second growth sites.

Pterostichus lama 2nd growth

0.4

0.35

h 0.3

0.25

0.2

0.15

0.1 Average seasonal catc seasonal Average

0.05

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

56 Figure 31. Mean seasonal catch (beetles per trap) of Pterostichus herculaneus (with associated standard error bars) plotted against patch area for old growth sites.

Pterostichus herculaneus old growth

1.00E+00

8.00E-01 h 6.00E-01

4.00E-01

2.00E-01

0.00E+00

Average seasonal catc seasonal Average 0 10000 20000 30000 40000 50000 60000

-2.00E-01

-4.00E-01 Patch area (sq. meters)

Figure 32. Mean seasonal catch (beetles per trap) of Pterostichus herculaneus (with associated standard error bars) plotted against patch area for second growth sites.

Pterostichus herculaneus 2nd growth

4.5

4

h 3.5

3

2.5

2

1.5

Average seasonal catc seasonal Average 1

0.5

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

57 Figure 33. Mean seasonal catch (beetles per trap) of Zacotus matthewsii (with associated standard error bars) plotted against patch area for old growth sites.

Zacotus matthewsii old growth

1.4

1.2

h 1

0.8

0.6

0.4

0.2 Average seasonal catc seasonal Average

0 0 10000 20000 30000 40000 50000 60000

-0.2 Patch area (sq. meters)

Figure 34. Mean seasonal catch (beetles per trap) of Zacotus matthewsii (with associated standard error bars) plotted against patch area for second growth sites.

Zacotus matthewsii 2nd growth

0.7

0.6 h 0.5

0.4

0.3

0.2 Average seasonal catc seasonal Average

0.1

0 0 2000 4000 6000 8000 10000 12000 Patch area (sq. meters)

58 Species Evenness

Whittaker plots of rank versus the log(percent abundance) were used to examine the dominance or egality of species within each community. Order for ranking was from most to least abundant. These plots were produced for each site for all dates pooled (Figures 35-37). I also examined the evenness among trapping locations within the group VR sites (Figures 38-46). There were more species and greater evenness in the large and medium group treatments of the VRAM experiment, than in the clearcut, old growth control or small patch treatments (Figure 35). Similarly, evenness and species richness were both greater in sites G4 and G5, than in the old growth control site from the VRAM (Figure 36). All the second growth group VR sites showed greater evenness and species richness than found in the 2nd growth control site. Group retention harvesting, in all cases, appeared to lead to a greater mix of species within a site, and greater evenness, whereas the control sites in both old growth and 2nd growth displaying the greater dominance of the forest specialists, particularly S. angusticollis, and fewer species overall. Differences among locations within sites were less apparent (Figures 38-46).

59 Figure 35. Whittaker plot for the old growth treatments of the VRAM experimental site.

Whittaker Plot Old Growth Sites

2

L 1.5 M Sm 1 C1 O1 0.5

0

Log (% abundance) (% Log -0.5

-1

-1.5 02468101214 Rank abundance

Figure 36. Whittaker plot for the old growth sites G4 and G5 together with the old growth control from the VRAM site.

Whittaker Plot Old Growth Sites

2

1.5 G4 G5 1 O1

0.5

0

Log (% abundance) (% Log -0.5

-1

-1.5 02468101214 Rank abundance

60 Figure 37. Whittaker plot for second growth sites including the 2nd growth control

Whittaker Plots 2nd growth sites

2 S1 G1 1.5 G2 G3 Large 1 G3 Small

0.5

0

Log abundance) (% -0.5

-1

-1.5 0 2 4 6 8 10121416 Rank abundance

Figure 38. Whittaker plot for all locations sampled within the group VR site G1.

Whittaker Plots G1 by location

2 G15to10mIn

1.5 G15to10mOut G1Edge G1In 1 G1Out

0.5

0

Log(%abundance) -0.5

-1

-1.5 02468101214 Rank abundance

61 Figure 39. Whittaker plot for all locations sampled within the group VR site G2.

Whittaker plots G2 by location

2

G25to10mOut 1.5 G2Edge G2In G2Out 1

0.5

0 Log(%abundance)

-0.5

-1 024681012 Rank Abundance

Figure 40. Whittaker plot for all locations sampled within the large patches of group VR site G3.

Whittaker plots G3 Large by location

2 G3Large5to10mIn G3Large5to10mOut 1.5 G3LargeEdge G3LargeIn 1 G3LargeOut

0.5

0 Log(%abundance)

-0.5

-1 0246810121416 Rank abundance

62 Figure 41. Whittaker plot for all locations sampled within the small patches within group VR site G3.

Whittaker plots G3 Small by location

2

G3Small5to10mOut G3SmallEdge 1.5 G3SmallIn G3SmallOut

1

0.5 Log(%abundance)

0

-0.5 02468101214 Rank abundance

Figure 42. Whittaker plot for all locations sampled within the group VR site G4.

Whittaker plots G4 by location 2 G45to10mOut G4Edge 1.5 G4In G4Out 1

0.5

0 Log(%abundance)

-0.5

-1 012345678910 Rank abundance

63 Figure 43. Whittaker plot for all locations sampled within the group VR site G5.

Whittaker plots G5 by location

2

G55to10mIn 1.5 G55to10mOut G5Edge G5In 1 G5Out

0.5

0 Log(%abundance)

-0.5

-1 024681012 Rank abundance

Figure 44. Whittaker plot for all locations sampled within the group VRAM site Sm.

Whittaker plots Sm by location 2

Sm5to10mOut SmEdge 1.5 SmIn SmOut

1

0.5 Log(%abundance)

0

-0.5 0123456789 Rank abundance

64 Figure 45. Whittaker plot for all locations sampled within the group VRAM site M.

Whitaker plots M by location

2 M5to10mOut MEdge 1.5 MIn MOut

1

0.5

0 Log(%abundance)

-0.5

-1 024681012 Rank abundance

Figure 46. Whittaker plot for all locations sampled within the group VRAM site L.

Whittaker plots L by location

2.5

L5to10mIn 2 L5to10mOut LEdge 1.5 LIn LOut 1

0.5

Log(%abundance) 0

-0.5

-1 012345678910 Rank abundnace

65 Heterogeneity measures The clearcut had the lowest species richness and species diversity overall. Among sites, there was greater species richness and diversity in the 2nd growth sites over the old growth sites (Table 18). Among 2nd growth sites, there was greater species richness and diversity in site G3, with a large number and variety of small and large patches, with lower diversity in sites G1 and G2 and the 2nd growth control. The old growth sites were similar in terms of diversity and richness- highest values overall were found in site G4, with a number of small patches. When I compared sites G1 (large patches) and G2 (small patches), it was apparent that there was greater species diversity and richness in the site G2 with smaller patches. Among the locations within each of these sites, there was apparently higher richness and diversity among In and Out locations for site G1, and for In and Edge locations for site G2 than found in the other locations (Table 19). When the large and small patches within site G3 were compared, it was apparent that species richness and diversity were greater in the small patches over the large patches. Among locations, there was greater species diversity and richness in Out locations around G3Large patches but no other differences were apparent (Table 20). For the comparison between small patch site G4 and large patch site G5, it was apparent that there was greater species richness, but lower species diversity in site G5 than in site G4. Within site G4, diversity was greater at the Edge and 5-10m Out from Edge locations than elsewhere in the site, but species richness was fairly even among locations (Table 21). Within site G5, diversity was generally even among locations, but species richness was higher in the In and Out locations than all others (Table 21). For the VRAM group treatments, it was apparent that highest species richness and diversity was found in the Large patch treatment, followed by the Medium patch treatment, with lowest values for both of these indices in the Small patch treatment (Tables 22&23). However, the differences in either of these indices among the three sites was minimal. Within the sites, species richness values were similar among locations, but species diversity was generally highest at the 5-10m In location in the Large treatment; there were higher values of species richness and diversity among the 5-10m Out from

66 Edge and Edge locations of the Medium treatment than in the other locations; and both values were generally even for all locations within the Small treatment (Tables 22&23).

67 Table 18. Species richness estimates and Diversity estimators (output from EstimateS, Colwell, 2004) C G1 G2 G3L G3S G4 G5 L M O S1 Sm ACE Mean 14.21 14.78 17.88 20.98 19.24 18.91 17.59 17.59 17.59 17.59 17.88 17.88 ICE Mean 33.67 24.5 18.74 20.58 18.63 17.99 17.42 17.45 17.44 17.46 17.53 17.52 Alpha Mean 1.91 1.99 2.22 2.29 2.22 2.17 2.12 2.1 2.07 2.06 2.02 2 Shannon Mean 1.39 1.67 1.88 1.79 1.78 1.73 1.7 1.68 1.67 1.67 1.69 1.71 Simpson Mean 3.25 3.69 4.64 4.02 3.92 3.68 3.52 3.42 3.44 3.43 3.56 3.65

Table 19. Species richness estimates and Diversity estimators by location within 2nd growth sites G1 (large patches) & G2 (small patches) G1 G1 G1 G1 G1 G2 G2 G2 G2 5to10mIn 5to10mOut Edge In Out 5to10mOut Edge In Out ACE Mean 11.31 12.52 13 13 13 15.11 15.11 14 ICE Mean 64.63 12.59 13.43 13 13 14.33 14.3 14 Alpha Mean 2.02 1.96 2.04 1.86 1.84 1.98 1.96 1.93 Shannon Mean 1.48 1.52 1.61 1.59 1.61 1.7 1.76 1.82 Simpson Mean 2.98 3.21 3.43 3.42 3.5 3.74 4 4.31

68 Table 20. Species richness estimates and Diversity estimators by location within 2nd growth sites G3 (large patches & small patches) G3Large G3Large G3Large G3Large G3Large G3Small G3Small G3Small 5to10mIn 5to10mOut Edge In Out Edge In Out ACE Mean 17.69 17.69 17.69 17.69 18.06 18.06 18.06 18.06 ICE Mean 17.44 17.51 17.56 17.62 17.6 17.6 17.6 17.6 Alpha Mean 2.11 2.09 2.09 2.09 2.06 2.06 2.05 2.05 Shannon Mean 1.69 1.7 1.71 1.72 1.71 1.71 1.72 1.72 Simpson Mean 3.44 3.51 3.56 3.6 3.58 3.6 3.64 3.65

Table 21. Species richness estimates and Diversity estimators by location within old growth sites G4 G4 G4 G4 G5 G5 G5 G5 G5 5to10mOut Edge In Out 5to10mIn 5to10mOut Edge In Out ACE Mean 15.55 15.55 15.55 15.55 17.88 18.11 18.36 22.4 22.4 ICE Mean 15.29 15.29 15.35 15.36 16.95 17.2 17.13 19.22 19.22 Alpha Mean 2.08 2.06 2.03 1.99 2.14 2.13 2.12 2.26 2.23 Shannon Mean 1.86 1.85 1.81 1.76 1.76 1.77 1.78 1.77 1.76 Simpson Mean 4.5 4.43 4.15 3.89 3.9 3.91 3.92 3.88 3.82

69 Table 22. Species richness estimates and Diversity estimators by location within the experimental VRAM old growth small and medium treatments. M M M M Sm Sm Sm SmOut 5to10mOut Edge In Out 5to10mOut Edge In ACE Mean 21.42 21.32 19.1 19.21 17.69 17.69 17.69 17.69 ICE Mean 19.06 19.23 18.14 18.09 17.44 17.44 17.44 17.44 Alpha Mean 2.18 2.17 2.16 2.13 2.13 2.12 2.12 2.11 Shannon Mean 1.72 1.72 1.7 1.68 1.68 1.69 1.69 1.68 Simpson Mean 3.6 3.6 3.52 3.42 3.45 3.46 3.48 3.42

Table 23. Species richness estimates and Diversity estimators by location within the experimental VRAM old growth large patch treatment. L L L L L 5to10mIn 5to10mOut Edge In Out ACE Mean 22.4 22.4 22.4 22.4 21.42 ICE Mean 19.22 19.24 19.24 19.24 19.06 Alpha Mean 2.22 2.21 2.2 2.2 2.18 Shannon Mean 1.76 1.73 1.73 1.73 1.71 Simpson Mean 3.83 3.67 3.67 3.69 3.59

70 Distance Coefficients

Seasonal catches by sites using weighted data The average catch of all species in the 2nd growth and old growth sites for all dates pooled was subjected to cluster analysis using average linkage to produce a similarity matrix based on Euclidean distance. Weighted averages were used for the group sites such that catches in any particular location were weighted by the % area that the particular location represented. Thus, in the Sm VRAM treatment, “In” catches were weighted by 5%, catches at the “Edge” were weighted as 19%, catches at the “5-10m Out” location were weighted as 24% and the “Out” catches were weighted as 52%. This weighting was related to the proposed area as estimated from setting maps (Table 24).

Table 24. Weighting of group sites by area occupied by edge, 5-10 m out from the edge, and 5-10 m in from the edge. VRAM In 5 to Out 5to10m Edge %retention 10m in out Old growth sites Sm 5 24 52 19 24 M 9 17 59 15 24 L 16 8 68 5.6 2.4 24 G4 3 28 53.6 15.4 18.4 G5 22 6 54 12 6 40

2nd growth sites G1 12 17.2 65 4 1.8 17.8 G2 7 22 51.7 19.3 26.3

G3Sm 4 15 64.2 16.8 20.8 G3L 14 15 64.2 3.8 3 20.8

The first cluster analysis, shown below, showed that when both old growth and 2nd growth sites were subjected to analysis, that the small patch sites G2 and Sm clustered together, that site G1 and G3 Small patches clustered together, and that the larger patch sites M, G5, L and G3 Large clustered closely together, with the small patch site G4 clustering somewhat separately.

71

Figure 47. Cluster analysis of all Group VR sites.

G1 G3SMALL G3LARGE M L G5 G4 SM G2

0.0 0.1 0.2 0.3 0.4 0.5 0.6 Distances

Next, Location Next, cluster analyses were performed on the data for average seasonal species catches by location. Firstly, cluster analysis was done for all old growth sites. It is apparent from Figure 48 that the catches from matrices or “Out” catches clustered separately from the catches from within patches or “In” catches. In addition, communities at “5 to 10m out from Edge” locations clustered with the matrix and clearcut communities, whereas communities of “5 to 10m in from Edge” locations clustered with the patch communities. Edge communities either cluster with matrix or patch communities, this varied by site. A similar demarcation is shown when cluster analysis was performed on locations within the VRAM treatments (Figure 49) and for sites G4 and G5 (Figure 50). In the latter case, the clearcut and old growth control VRAM treatments were included for comparison with these two old growth sites. All the “5-10m out from Edge”, and matrix communities clustered with the clearcut, whereas the patch communities, “5-10m in from Edge” and edge communities clustered with the old growth control.

72 Next, cluster analysis was done for 2nd growth sites. Firstly, clustering was done for all locations within all 2nd growth sites (G1, G2, G3S, G3L and the 2nd growth control) (Figure 51). The catches from the small patches of site G2 (from both the edge and within patches) clustered with catches from matrices of other sites. In addition, there was close clustering of catches from the edge and patches of the small patches of site G3 together with matrix (“Out”) catches. Conversely, catches from within large patches and edges of sites G3 and G1 clustered together with the 2nd growth control, as well as catches from “5-10m in from the edge” locations in site G1. When we examine just the communities from the small and large patches of site G3 together with the 2nd growth control, it is apparent that catches from within large patches, and 5 to 10m in from the edges of patches in site G3 cluster with the 2nd growth control, but catches from all locations around small patches of site G3 clustered with the communities of the edge, 5 to 10 m out from the edge, and matrix around large patches in site G3 (Figure 52). When we look at the communities of sites G1 (large patches) and G2 (small patches), it is apparent that the catches from within patches, as well as edges of patches and 5-10m out from the edge of patches of site G1 cluster together with the 2nd growth control, whereas catches from all locations, even within the small patches, of site G2, cluster with the matrix catches of site G1 (Figure 53).

73 Figure 48. Cluster analysis of seasonal carabid catches by location within old growth sites and VRAM treatments.

C1 G4OUT SMOUT LOUT SM5TO10MOUT MOUT LEDGE L5TO10MOUT M5TO10MOUT G45TO10MOUT G5OUT SMEDGE G55TO10MOUT G55TO10MIN SMIN MEDGE O1 G4EDGE MIN G4IN G5EDGE G5IN LIN L5TO10MIN 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Distances

Figure 49. Cluster analysis of seasonal carabid catches by location within the VRAM treatments (clearcut, old growth, small, medium and large group retention

patches).

C1 SMOUT LOUT SM5TO10MOUT MOUT LEDGE L5TO10MOUT M5TO10MOUT SMEDGE SMIN MEDGE O1 MIN LIN L5TO10MIN

0.0 0.5 1.0 1.5 Distances

74 Figure 50. Cluster analysis of seasonal carabid catches by location within old growth sites G4 and G5, together with the Clearcut and Old Growth control catches from the VRAM treatments.

C1 G4OUT G5OUT G45TO10MOUT G55TO10MOUT G55TO10MIN G4IN G5EDGE O1 G4EDGE G5IN

0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 Distances

Figure 51. Cluster analysis of seasonal carabid catches by location within the 2nd nd growth sites (G1, G2, G3Large, G3Small and 2 growth control).

G35TO10MINLA S1 G3INLARGE G1IN G15TO10MIN G1EDGE G15TO10MOUT G35TO10MOUTS G1OUT G3EDGESMALL G3INSMALL G25TO10MOUT G2IN G3EDGELARGE G2EDGE G2OUT G3OUTLARGE G35TO10MOUTL G3OUTSMALL

0.0 0.5 1.0 1.5 Distances

75 Figure 52. Cluster analysis of seasonal carabid catches by location within the 2nd nd growth sites G3Large and G3Small together with the 2 growth control.

G3INLARGE S1 G35TO10MINLA G3INSMALL G3EDGESMALL G3OUTSMALL G3EDGELARGE G3OUTLARGE G35TO10MOUTL G35TO10MOUTS

0.0 0.5 1.0 1.5 Distances

Figure 53. Cluster analysis of seasonal carabid catches by location within the 2nd nd growth sites G1 and G2 together with the 2 growth control.

G1IN G15TO10MOUT G1EDGE G15TO10MIN S1 G1OUT G25TO10MOUT G2IN G2EDGE G2OUT

0.0 0.5 1.0 1.5 Distances

76 DISCUSSION

Currently, there is a great deal of concern about the impact of habitat loss and fragmentation on biodiversity (Saunders et al. 1987; Doherty & Grub 2002). Studies that document the decline of species from habitat fragments suggest that about one-fifth to one-third of species decline and this has been noted in a wide range of environments, landscape contexts and among different species e.g. (Klein 1989, Didham et al. 1998)[beetles]; Saunders 1989[birds]; Rosenblatt et al. 1999[mammals]; Chamberlain et al. 2000[birds]; Nupp & Swihart 2000[mammals]; Driscoll 2004[reptiles]). Beetles appear to suffer a similar magnitude of decline as vertebrates in fragmented landscapes (Klein 1989; Saunders 1989; Rosenblatt et al. 1999; Driscoll 2004). However, there have been a few exceptions, such as low number of declining beetles from an agricultural landscape in Finland (Halme & Niemela 1993), possibly due to a relatively recent glaciation, and in one other case, where only 9% of beetle species declined and were absent from the matrix, possibly due to the very short period since the fragmentation event (Davies et al. 2000). Fahrig (1997) has suggested that landscapes cleared to levels below 20% cross a threshold below which species suffer a rapidly increasing risk of extinction.

Some species of beetles are apparently particularly area sensitive and thus become extinct in patches that are too small (Bolger et al. 2000, Lindenmayer et al. 2002), but others can apparently survive well in small patches. For example, in Amazonian forest fragments, 62% of beetle species are expected to survive in a landscape of many 1-ha fragments and 90% of species are expected to survive in a landscape of many 100-ha fragments (Didham et al. 1998). Some species survive because they can become abundant in the matrices surrounding habitat patches, and are thus able to alter species composition and increase species richness in remnant vegetation (Webb & Hopkins 1984; Halme and Niemela 1993). My work in previous years showed that many species were indeed able to survive in patches of different sizes, particularly generalist species that did not appear to depend on forested areas for their habitat (Pearsall 2002, 2003).

77 The reasons for species decline after habitat fragmentation may be varied. Relationships between response and species characteristics such as trophic level (Komonen et al. 2000), body size (Cosson et al. 1999), and dispersal ability (Thomas 2000) have been examined. It is generally believed that species with good dispersal ability are more likely to survive fragmentation by avoiding local extinctions through a rescue effect (Brown & Kodric-Brown 1977) or by being able to recolonize vacant habitat patches after local extinction (Hanski & Gilpin 1997). Driscoll and Weir (2005) found that the dispersal capacity of beetles was a major predictor of how they responded to fragmentation in woodlands in Australia. Many studies have shown a negative relationship between isolation and both species richness and abundance in fragments, which has given further credence to the idea that dispersal ability influences species' responses to fragmentation (e.g., Davies et al. 2000). Several empirical studies have also highlighted the importance of high dispersal as a factor promoting survival in fragmented landscapes (e.g., de Vries et al. 1996; Thomas 2000), yet conversely, results from other studies have shown no effect of dispersal ability on survival in fragments (e.g. MacNally et al. 2000).

My work during previous years has suggested that although patches in group retention harvesting may be effective at preservation of forest specialists, most of which are flightless, and with low dispersal ability, in the very short term (ie one-year post- harvest), that this may not be borne out in older VR sites, particularly those with small patches or low levels of retention. Work done in 2003 in remnant patches of old growth in North Vancouver Island, that had been in existence for longer periods of time (35 years +), did show some differences in carabid assemblages to uncut tracts of old growth forest, which also suggested some longer term responses to fragmentation (Pearsall, 2004).

During 2005, I began a detailed examination of the effect of patch size on carabid communities in both experimental and operational forest blocks in West Island Timberlands. In particular, I was interested in comparing how well patches of different size retain original forest communities of beetles, as well as to examine edge effects, the potential “lifeboating” of patches, and how the size of patches affects composition of

78 beetle communities within the cut matrices. The work was carried out in both 2nd growth and old growth sites, so I was also interested in examining whether the findings were consistent among old growth and 2nd growth sites.

A total of three group retention sites (G1, G2, G3) were chosen in 2nd growth forest, and two group retention sites (G4 & G5) were chosen in old growth forest in West Island Timberlands. Site G1 contained large patches whereas those in site G2 were small. Both small and large patches were found in site G3. Site G4 had small patches and site G5 had two large patches. Traps were also set in 2nd growth forest located adjacent to site G2 as our 2nd growth control site (S1). I also set traps in the Klanawa group size VRAM site, which consisted of 5 treatments: old growth control (O), small patches (Sm), medium size patches (M), large patches (L) and clearcut ( C). Medium sized patches in the VRAM site were a similar size to the patches in site G4. All four 2nd growth sites were in the CWHxm and all old growth sites were in the CWHvm1. All sites had been harvested in 2003/2004 except for the VRAM site which was harvested in 2005. All 3 VRAM treatments (small patches, medium patches and large patches) had the same retention level of 24.3%. All the other sites had similar retention levels around 18-25% with the exception of site G5, where retention levels were 39.9%.

A total of 291 traps were placed out in the operational (G1-G5, S1) and experimental sites (old growth and medium sized patches) in late April, and a further 86 were placed out in the experimental site (small patches, clearcut and large patch treatments) in late May. Collections were made every month for May-September inclusive.

Prior work has identified Scaphinotus angusticollis, Pterostichus crenicollis and Zacotus mathewsii as forest species, and P. lama and P. herculaneus as generalist species, and thus I focus most of the discussion on patterns that I saw in this group of species, which were also the most common species caught (Pearsall 2002).

79 The 2nd growth sites had generally greater species richness and diversity than the old growth sites, and control sites in both forest types generally had lower species richness and diversity than the group retention sites, probably because of influx of new species associated with new habitats and forest edges that were provided by group retention harvesting. Some species were only found in second growth forest, such as Promognathus crassus, and Omus dejeani, whereas Pterostichus pumilus pumilus was only caught in old growth sites. Species richness did not vary greatly between small and large patch sites in 2nd growth forest, but was much lower in the small patch site G4 than the large patch site G5 in the old growth, and much lower in the small patch site (Sm) of the VRAM experiment than in the other treatments. Species diversity was generally greater in small patches over large patches except for in the VRAM site, where diversity among trreatments was generally similar, but did show a slight declined as patch size declined. The difference may be due to the fact that this experimental site had been very recently logged, whereas the other sites had had a few years to recover. Consistent patterns in species richness and diversity among locations could not be discerned, although several sites did show an increase in diversity at or close to the edge. There was particularly low species richness and species diversity in the clearcut treatment, possibly because many of the forest species were unable to survive in this site, but populations of disturbance specialists and generalists may have been slow to repopulate this site, which was heli- logged and quite isolated. In terms of species abundance, more beetles per trap were captured in the 2nd growth sites than the old growth sites. Within those sites, there was greater mean abundance in the 2nd growth control, with lower abundance in the large patch sites, and lowest abundance overall in the small patch sites. Within the old growth sites, mean abundance was highest in the large patch sites and the old growth control, and progressively lower in the sites as patch size declined, with lowest catches overall in the clearcut. This drop in overall abundance is associated with the decline in the forest specialist, Scaphinotus angusticollis.

80 Seasonal patterns of abundance showed general peaks of abundance in May and September in most sites, for both total carabids pooled and for Scaphinotus angusticollis, the most commonly captured beetle, which is the general pattern shown in previous studies (Pearsall, 2003, 2004, 2005). These months coincide with reproductive periods, and generally, both activity and abundance are greatest at this time, which suggests that on a more limited budget, that these would be the optimal trapping periods, at least on Vancouver Island.

Patches in all sites were able to retain forest specialists, and abundance of forest specialists showed a positive relationship to size of patch. Large patches always contained higher numbers of forest specialists per trap than small patches. This was the case for both old growth and 2nd growth sites. Retention of forest specialists was generally better in old growth patches than 2nd growth patches, and small patches of 2nd growth sites were particularly poor at retention of species such as S. angusticollis, Z. matthewsii, and P. crenicollis.

Forest specialists increased in abundance at the patch-matrix transition, with highest catches inside of patches. Some species showed an accumulation close to or at the edge, for example, Cychrus tuberculatus, P. crassus, P. crenicollis, P. herculaneus, Z. matthewsii. This response by Z. matthewsii has been noted in prior studies (Pearsall, 2003). The forest specialist species (S. angusticollis, P. crenicollis and Z. matthewsii) showed an expected increase inside of patches as compared to in the matrices, while generalist species such as P. lama, P. crassus, C. tuberculatus and P. herculaneus generally showed no clear differences in abundance among patch and matrix locations. Forest specialists were no more abundant in the matrices of the VRAM group retention treatments than in the clearcut, so in this site, life-boating was not apparently occurring. However, the VRAM had only recently been logged, and activity was still occurring in a number of the sites as we began our work. Indeed, this led to a loss of a number of the traps in the Sm site in June, due to their destruction. However, it is possible that there had not been enough time in this site for the vegetation to recover in

81 the matrices, and thus forest specialists were not able to venture into the dry and vegetation- bereft matrix surrounding patches.

Cluster analysis showed that communities of the patches and matrices of the old growth sites were very dissimilar, and that carabid beetles showed immediate responses at the forest edge, such that communities at locations 5 to 10 m into the matrix from the edge of patches, were similar to the matrix communities, whereas communities just 5 to 10 m in from the patch edge were similar to the patch communities. In all cases, patch communities were more similar to the old growth control communities, whereas matrix communities were more similar to the clearcut community. For second growth sites, a similar relationship existed, except for these sites, there was also a clear demarcation among the communities of the small versus large patches. The communities of the small patch sites, G2 and G3S, were more similar overall to matrix communities than to the 2nd growth control, whereas the large patch site communities were more similar to the 2nd growth control. As noted earlier, forest specialists were less well retained in the small patches in 2nd growth sites, and this pattern is supported by the cluster analysis results.

Using these data, we were able to put together species-patch area curves and examine the direct response of a number of the forest specialists to patch size. My results showed that there was a clear response by these species (S. angusticollis, P. crenicollis and Z. mathewsii) to patch size, with increasing catches per trap in larger patches. However, the larger catches in larger patches did not appear to be related to increased life-boating ability of larger patches, as forest specialists were no more abundant in the matrices surrounding any of the patches than in the clearcut. This may be related to the age of the harvested matrices, and future comparisons might show that repopulation of the matrices occurs quicker than expected over time than would take place in the traditional clearcut.

How do these results compare with those of other studies of the effects of forest fragmentation on beetles? Other studies carried out world-wide have examined the

82 response of carabids to forest harvesting, to patch retention harvesting, and have examined the distribution of carabid assemblages across forest edges (e.g. Heliola et al. 2000, Lemieux and Lindgren 2004, Niemela 1997). Some current studies in B.C include an examination of carabid communities of chronosequences in the Houston Forest Products area near Houston and Burns Lake, focussing primarily on the relationship between species and CWD (Staffan Lindgren, UNBC, pers. comm.), and an examination of the ecological responses of a carabid species, Scaphinotus angusticollis to forest fragments left in riparian areas of the Malcolm Knapp Research Forest. Didham et al. (1998) examined the effects of forest fragmentation on leaf-litter beetle species in an experimentally fragmented tropical forest in Central Amazonia. They found that most of the species were adversely affected, with species loss rates increasing with decreasing patch size. In this case, declining densities were noted prior to species loss from smaller patches- my findings of decreasing densities of a number of forest specialists during this study may also be a warning flag for the potential for future species loss from the smaller patches examined. During 2004, my work in some of the oldest operational group VR sites in West Vancouver Island showed that the group retention sites were very bereft in terms of species richness, and that mean catches of forest specialists in the patches were far lower than found in control sites. My conclusion was that these operational VR sites were not able (4 years post-harvest) to retain the original forest communities adequately. I had noted that 2004 had been a hot and dry year and that possibly this might have resulted in reduced survivorship in these sites- however, the communities of the control sites did not appear to be equally affected, suiggesting that this was a VR treatment and patch size effect rather than a climate response. It will be important to examine some of the older VR sites again in wetter years in the future, especially those where levels of retention or patch size is low, to determine whether the loss of species richness and diversity noted in 2004 was an anomaly related to weather conditions, or whether the patches were truly too small to retain communities adequately.

Niemela et al. (1993) and Niemela (1996) noted that logged areas generally had greater species richness and more heterogeneous communities than undisturbed areas. They noted that such areas were in a state of flux with regard to beetle communities,

83 which were often typified by influx of open-habitat species such as Amara and Harpalus, with their characteristic high population turnover rates. My results of increasing species richness and diversity in group retention sites over that found in the control sites and old growth sites would be expected, given the loss in overall abundance of forest specialists such as S. angusticollis and the opening up of new habitats and edges for repopulation. Disturbance species such as Notiophilus sylvaticus, Amara idahoensis, and Harpalus animosus were only found in group VR site, and were not found within the control sites in the study this year. Spence et al (1996) noted that open-habitat specialists were able to colonize forest patches up to 80 m from the forest edge, and that the ability of patches to retain forest specialists would probably decrease over time, if the open-habitat species were able to compete with the forest specialists. In my study, few of the sites had patches this size, except for the large patches of site G5, L of the VRAM and patches in site G1, and thus we might have expected to find the reduced abundance of forest specialists that was apparent in patches less than 80 m diameter (ie. Sites G2, G3S, G4, M and Sm). Interestingly, Z. matthewsii, which has been noted as an old-growth specialist in prior studies (at the HJ Andrews Experimental Forest in Oregon and by Craig (1995) for southern Vancouver Island), was only well retained in patches of site G5 over all other sites: the patches in this site were largest of all the patches sampled during the course of the study, and % retention was also greatest in this site. This may be an important red flag species, since is shows such a strong response to patch size as well as its build-up at the edge of sites, which has been noted in some of my previous studies (Pearsall, 2002).

Ghandi et al (2003) found patch shape to be an important determinant of carabid species retention when they examined harvest residuals in coniferous forests of northern Alberta, but interestingly, noted that the same response was not seen for staphylinid beetles. Specifically, they found that round harvest residuals contained greater numbers of carabid individuals, and showed greater aggregation of forest specialists than elliptical patches, which they suggested was due to the difference in distance to edge in the differently shaped patches. This led them to postulate that the retention of subtle interior forest characteristics might be critical for retention of forest specialists, and that this might be better achieved with a circular shape. This factor was not examined in this

84 study, but it may be noted that patches, although primarily circular in shape, were sometimes long and thin due to blowdown events. Although I did not work in many patches that were of this shape, it is a factor that could be considered in future studies.

Osawa et al. (2005) examined carabid beetles before and after logging in an urban forest in central Japan. They examined specific microclimatic and vegetation responses prior to and post logging, and concluded that abundance of beetles was directly related to vegetation diversity and thus source of food, prior to harvesting. After harvesting, however, this factor was not significant, probably because logging altered the habitat characteristics and food availability for the beetles. The authors suggested that large generalist carnivores and insectivores appeared more vulnerable to the environmental changes created after logging, than were small species of beetles. Our forest specialists, Z. matthewsii, and S. angusticollis are both carnivores, the former feeding on millipedes, and the latter on slugs and snails, and both are large, and cannot fly out of patches.

In conclusion, my results show a strong response by carabid beetles to forest edges and patch sizes. 3 years post-harvest, patches were able to retain forest specialists in all of the sites, but carabid beetles showed a response to patch size in group retention sites, with higher catches per trap of forest specialists in larger patches than in smaller patches, a response noted in both old growth and 2nd growth sites. An important finding was that retention of forest specialists was generally better in old growth patches than 2nd growth patches, and small patches of 2nd growth sites were particularly poor at retaining species such as S. angusticollis, Z. matthewsii, and P. crenicollis. These results may have important implications for patch size considerations in differently aged forests of Vancouver Island. Future studies should examine old growth in CWHxm and 2nd growth in CWHvm1, so that we can directly compare 2nd growth and old growth responses to patch size within similar variants.

85 CONCLUSIONS

1. 3 years post-harvest, VR patches were able to retain forest specialists in all of the sites. However, carabid beetles showed a response to patch size in group retention sites, with higher catches per trap of forest specialists in larger patches than in smaller patches. This response was seen in both old growth and 2nd growth sites. However, retention of forest specialists was generally better in the old growth patches than the 2nd growth patches, and small patches of 2nd growth sites were particularly poor at retaining species such as S. angusticollis, Z. matthewsii, and P. crenicollis.

2. Carabid beetles showed edge responses, and several species built up at or close to the edge of patches. This may be associated with a general unwillingness by forest specialists to venture further out into the matrix. However, some of the generalist species also built up at the edges, which suggests that this may be related to the change in vegetation or microclimate at this location.

3. The 2nd growth sites had generally greater species richness and diversity than the old growth sites, and control sites in both forest types generally had lower species richness and diversity than the group retention sites, probably because of influx of new species associated with new habitats and forest edges that were provided by group retention harvesting. Diversity was generally greater in small patch sites than sites with larger patches, except in the VRAM site, where diversity indices were very similar among the group retention treatments. Higher diversity in the small patch sites may be associated with the greater amount of edge, and greater size of matrix tracts, which may influence forest specialists in a negative fashion, and allow for greater abundance of forest generalists.

4. My results show a strong response by carabid beetles to forest edges and patch sizes. Whether this is a result of the microclimate or prey base of small patches being less ideal than larger patches, or an isolation effect, remains to be determined.

5. This year we had two replicate sites for both large patches and small patches in each of the old growth and 2nd growth sites. It will be important to try to add another replicate pair (small and large patch sites) in 2nd growth in CWHxm and another replicate pair of sites for old growth in the CWHvm1. If possible, it would be useful to also add 3 replicate pairs of large and small patch sites in the 2nd growth in CWHvm1 so that we can compare old growth and second growth sites in the same variant.

86 ACKNOWLEDGEMENTS

I am very grateful to Bill Beese for his help and assistance with this project and to Jeff Sandford for invaluable help with location of blocks, loans of field equipment and advice and help with data and mapping needs. This work could not have been done without the great field assistance of Claudia Lake. Thank you very much to Jeff Jarrett for help with beetle identifications, to Karen Needham for allowing me space and time in the UBC George J. Spencer Entomological Museum. Thanks to the staff at West Island Timberlands, particularly Jon Flintoft and Mike Davis, with their assistance and advice regarding block locations and information.

87

REFERENCES

Bolger, D. T., A. V. Suarez, K. R. Crooks, S. A. Morrison, and T. J. Case. 2000. Arthropods in urban habitat fragments in southern California: area, age, and edge effects. Ecological Applications 10:1230–1248.

Brown, J. H., and A. Kodric-Brown. 1977. Turnover rates in insular biogeography: effect of immigration on extinction. Ecology 58:445–449.

Burke, D., & H. Goulet 1998. Landscape and area effects on beetles assemblages in Ontario. Ecography 21: 472-479

Chamberlain, D. E., R. J. Fuller, R. G. H. Bunce, J. C. Duckworth, and M. Shrubb. 2000. Changes in the abundance of farmland birds in relation to the timing of agricultural intensification in England and Wales. Journal of Applied Ecology 37:771– 788.

Clarke, K.R. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18: 117-143

Colwell, R. K. 2004. EstimateS: Statistical estimation of species richness and shared species from samples. Version 7. User's Guide and application published at: http://purl.oclc.org/estimates

Cosson, J. F., S. Ringuet, O. Claessens, J. C. de Massary, A. Dalecky, J. F. Villiers, L. Granjon, and J. M. Pons. 1999. Ecological changes in recent land-bridge islands in French Guiana, with emphasis on vertebrate communities. Biological Conservation 91:213–222.

Craig, K.G. (1995) Variation in carabid community structure associated with coastal Douglas-Fir forest successional stages. M.Sc. Thesis, University of British Columbia.

Davies, K. F., C. R. Margules, and J. F. Lawrence. 2000. Which traits of species predict population declines in experimental forest fragments? Ecology 81:1450–1461.

de Vries, H. H., P. J. denBoer, and T. S. vanDijk. 1996. Ground beetle species in heathland fragments in relation to survival, dispersal, and habitat preference. Oecologia 107:332–342.

Didham, R. K., J. Ghazoul, N. Stork, and A. J. Davis. 1996. Insects in fragmented forests: a functional approach. Trends in Ecology & Evolution 11:255–260.

88 Didham, R. K., P. M. Hammond, J. H. Lawton, P. Eggleton, and N. E. Stork. 1998. Beetle species responses to tropical forest fragmentation. Ecological Monographs 68:295–323.

Doherty, P. F., and T. C. Grubb. 2002. Survivorship of permanent-resident birds in a fragmented forested landscape. Ecology 83:844–857.

Driscoll, D. A. 2004. Extinction and outbreaks accompany fragmentation of a reptile community. Ecological Applications 14:220–240.

Driscoll, D. A., and T. Weir. 2005. Beetle Responses to Habitat Fragmentation Depend on Ecological Traits, Habitat condition, and Remnant Size. Conservation Biology 19: 182-202

Fahrig, L. 1997. Relative effects of habitat loss and fragmentation on population extinction. Journal of Wildlife Management 61:603–610.

Gandhi, K.J.K., J.R. Spence, D.W. Langor, L.E. Morgantini, and K.J. Cryer. 2004. Harvest Retention patches are insufficient as stand analogues of fire residuals for litter dwelling beetles in northern coniferous forests. Canadian Journal of Forest Research 34: 1319-1331

Haila, Y., I.K. Hanski, J. Niemela, P. Puntilla, S. Raivio, & H. Tukia 1994. Forestry and boreal fauna: matching management with nautral forest dynamics. Annales Zoologici Fennici 30: 17-30

Halme, E., and J. Niemela. 1993. Carabid beetles in fragments of coniferous forest. Annales Zoologici Fennici 30:17–30.

Hanski, I., and M. E. Gilpin. 1997. Metapopulation biology. Ecology, genetics and evolution. Academic Press, San Diego.

Harris, L.D. 1984. The fragmented forest: island biogeography theory and the preservation of biotic diversity. University of Chicago Press, Chicago.

Heliola, J, M. Koivula, & J. Niemela 2000. Distribution of carabid beetles (Coleoptera, Carabidae) across a Boreal Forest-Clearcut Ecotone. Conservation Biology: 370-377

Klein, B. C. 1989. Effects of forest fragmentation on dung and carrion beetle communities in central Amazonia. Ecology 70:1715–1725.

Komonen, A., R. Penttilä, M. Lindgren, and I. Hanski. 2000. Forest fragmentation truncates a food chain based on an old-growth forest bracket fungus. Oikos 90:119–126.

89 Krebs, C.J. 1989. Ecological Methodology. Harper and Row Publishers, New York. 654pp.

Lemieux, J.P. & Staffan Lindgren, B. 2004. Ground beetle responses to patch retention harvesting in high elevation forests in B.C. Ecography 29: 557-566

Lindenmayer, D. B., R. B. Cunningham, C. F. Donnelly, H. Nix, and B. D. Lindenmayer. 2002. Effects of forest fragmentation on bird assemblages in a novel landscape context. Ecological Monographs 72:1–18.

Lindroth, C. H. 1961-1969. The ground beetles (Carabidae excl. Cicindelinae) of Canada and Alaska. Parts 1-6. Opuscula Entomologica xlviii + 1192 pp.

MacNally, R., A. F. Bennett, and G. Horrocks. 2000. Forecasting the impacts of habitat fragmentation. Evaluation of species-specific predictions of the impact of habitat fragmentation on birds in the box–ironbark forests of central Victoria, Australia. Biological Conservation 95:7–29.

Magurran, A.E. 1988. Ecological Diversity and its Measurement. Princeton University Press, Princeton, N.J.

Matlack, G.R.& J.A. Litvaitis 1999. Forest edges. Pages 210-233 in M.L. Hunter, Jr., editor. Maintaining biodiversity in forest ecosystems. Cambridge University Press, Cambridge, U.K.

Niemelä, J., Langor, D. & Spence, J. R. 1993: Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae) in western Canada . Conservation Biology 7: 551-561.

Niemelä, J. (editor). 1996: Population biology and conservation of carabid beetles. Annales Zoologici Fennici 33:1-241.

Niemela, J. 1997. Invertebrates and boreal forest management. Conservation Biology 11: 601-610

Nupp, T. E., and R. K. Swihart. 2000. Landscape-level correlates of small- mammal assemblages in forest fragments of farmland. Journal of Mammalogy 81:512– 526.

Osawa, N., A. Terai, K. Hirata, and A. Nakanishi. 2005. Beetle Responses to Habitat Fragmentation Depend on Ecological Traits, Habitat Condition, and Remnant Size. Canadian Journal of Forest Research 35(11):2698-2708

90 Pearsall, I.A. 2000. Carabid beetles (Coleoptera: Carabidae): Are these good indicator species to be used in an adaptive management program and how might we carry out a pilot assessment? Contractual report to Weyerhaeuser December 2000.

Pearsall, I.A. 2002. Study to Assess the Efficacy of Ground Beetles (Coleoptera: Carabidae) as Ecological Indicators. Final report to Weyerhaeuser March 2002.

Pearsall, I.A. 2003. Study to Assess the Efficacy of Ground Beetles (Coleoptera: Carabidae) as Ecological Indicators In Two Variable-Retention Experimental Sites. Final report to Weyerhaeuser March 2003.

Rosenblatt, D. L., E. J. Heske, S. L. Nelson, D. H. Barber, M. A. Miller, and B. MacAllister. 1999. Forest fragments in east-central Illinois: islands or habitat patches for mammals? American Midland Naturalist 141:115–123.

Saunders, D. A. 1989. Changes in the avifauna of a region, district and remnant as a result of fragmentation of native vegetation—the wheatbelt of Western Australia—a case study. Biological Conservation 50:99–135.

Southwood, T.R.E. 1978. Ecological methods: with special reference to the study of insect populations. Chapman and Hall, New York.

Spence, J.R., D. Langor, J. Niemela, H. Caracamo & C. Currie 1996. Northern forestry and carabids: the case for concern about old-growth species. Annales Zoologici Fennici 33: 173-184

Stevens, S.M. & T.P. Husband 1998. The influence of edge on small mammals: evidence from Brazilian Atlantic forest fragments. Biological Conservation 85: 1-8

Thomas, C. D. 2000. Dispersal and extinction in fragmented landscapes. Proceedings of the Royal Society of London Series B-Biological Sciences 267:139–145.

Von Sacken, A. 1998. Interior habitat. Pages 130-145 in J. Voller & S. Harrison, editors. Conservation Biology Principles for forested landscapes. UBC Press, Vancouver, Canada.

Webb, N. R., and P. J. Hopkins. 1984. Invertebrate diversity on fragmented Calluna heathland. Journal of Applied Ecology 21:921–933.

Zar, J.H. 1984. Biostatistical Analysis. Prentice-Hall.

91 APPENDIX ONE: SITE CHARACTERISTICS 2005. Pre-harvest data obtained from VR-mdb (customized by Jeff Sandford) for the experimental VRAM site Clearcut Old growth Small patch Medium patch Large patch % coverage 82.4 73.3 75 68.3 69.5 (by deciduous and coniferous trees) pre- harvest % moss cover 8.6 14.7 16.7 16.7 15.3 % shrub cover 9.5 31.7 35.3 35.3 27.5 % herb cover 24.2 20.9 11.4 11.4 22.1 % litter cover 82.9 88.5 86.7 86.7 88.1 Most GAULSHA, GAULSHA, GAULSHA, GAULSHA, GAULSHA, commonly VACCPAR, VACCPAR, VACCPAR, VACCPAR, VACCPAR, recorded RUBUSPE VACCALA, RUBUSPE, RUBUSPE, RUBUSPE, shrub species RUBUSPE VACCALA, VACCALA, MENZFER MENZFER Most POLYMUN, BLECSPI, POLYMUN, BLECSPI, POLYMUN, commonly BLECSPI, POLYMUN, BLECSPI, POLYMUN, BLECSPI, recorded herb TRILOVA TRILOVA, TIARTRI, TIARTRI, TRILOVA, species TIARTRI TRILOVA TRILOVA, TIARTRI, LYSIAME Shrub species: VACCPAR (Red huckleberry), RUBUPAR (Thimbleberry), MAHONER (Dull Oregon-grape), GAULSHA (Salal), ROSAGYM (Baldhip rose), MENZFER (False Azalea), VACCOVL (Oval leaved blueberry), RUBUURS (Trailing blackberry), RUBUSPE (Salmonberry), VACCALA (Alaskan blueberry)

92 Herb species: POLYMUN (Sword fern), TIARTRI (Three-leaved foam flower), LYSIAME (Skunk-cabbage), BLECSPI (Deer fern), LINNBOR (Twinflower), PTERAQU (Bracken fern), ATHYFIL (Lady fern), EPILANG (fireweed), ACHLTRI (Vanilla leaf), ANAPMAR (Pearly Everlasting), TRILOVA (Western Trillium)

Post-harvest data for 2nd growth sites (using same codes as above) G1 G2 G3 S1 IN Various mosses, VACCPAR, various mosses, POLYMUN, lot TIARTRI, POLYMUN, ACHLTRI, so mosses & Rubus sp., GAULSHA, GAULSHA, liverworts, RUBUSPE, various mosses, POLYMUN, VACCPAR, POLYMUN, VACCPAR, ACHLTRI, GAULSHA, POLYMUN GAULSHA OUT RUBUSPE, PTERAQU, PTERAQU, POLYMUN, Rubus sp., EPILANG, GAULSHA, BLECSPI, GAULSHA ANAPMAR, GAULSHA, Rubus sp., EPILANG, POLYMUN, RUBUSPE, Rubus sp., RUBUSPE, RUBUPAR, VACCPAR ANAPMAR, POLYMUN, PTERAQU, VACCPAR. ACHLTRI, BLECSPI EPILANG, VACCOVL

93 APPENDIX TWO: PHOTOGRAPHS OF SITES

Site G1 large patches

Edge of a large patch in site G1

94 Site G2 small patches and the surrounding matrix

Site G2 matrix

95 Site G3 Large Patches

Inside a large patch at site G3

96

Matrix outside of a small patch at site G3

Edge of a small patch at site G3

97

2nd growth control (S1)

Trap in the 2nd growth control site

98 Site G4 Small patches

Site G4 small patches

99

Site G5 Large patches

Inside a patch at site G5

100

Matrix surrounding site G5

101 Large patch VRAM treatment

Edge of one of the large patches at the VRAM site

102 Within patches of Large patch VRAM treatment

Medium patch VRAM treatment

103 Medium patch VRAM treatment

Inside a medium sized patch at the VRAM site

104 Small patch VRAM treatment

Inside a small patch VRAM treatment

105 Clearcut VRAM treatment

View of the clearcut (heli-logged) treatment from the road entrance

106

View from the middle of the clearcut site down to the Klanawa valley below.

Old growth site VRAM

107

APPENDIX THREE: NEW SPECIES DISCOVERED DURING 2005

Pearsall (2002) lists species descriptions and biological information for the following species: Scaphinotus angusticollis, S. marginatus, S. angulatus, Notiophilus sylvaticus, Pterostichus lama, P. herculaneus, P. amethystinus, P.adstrictus, P. castaneus, Bembidion irridescens, Zacotus matthewsii, and Leistus ferruginosus. Pearsall (2003) species descriptions and biological information for the following species: Loricera decempuntata, Synuchus impunctatus, Amara littoralis, Amara obesa, Harpalus somnulentus, Bembidion approximatum and Cychrus tuberculatus. Pearsall (2004) lists species descriptions and biological information for the following species: Carabus granulatus, C. nemoralis, C. taedatus, Harpalus cordifer, H. cautus, Promognathus crassus and Omus dejeani. Pearsall (2005) lists species descriptions and biological information for the following species: Amara erratica and Pterostichus pumilus pumilus.

Two species were captured this year which had not been caught in previous years of the study. Very little information was available for either of them. Both are very small, winged species, and were only found as single specimens during the study.

Harpalus animosus Casey 1924 Lindroth stated that this species was found on alpine and sub-alpine meadows, sometimes appearing immediately after snow-melt.

Amara idahoana Casey 1924 This species was found in open “parks” above 5500ft (Minks & Hch. 1939, in Lindroth 1946). Most individuals apparently possess fully developed wings but in some, the wings were narrower than one elytron and believed to be non-functionary.

108 APPENDIX FOUR: ANALYSIS OF BY-CATCH

By-catch during 2005 consisted of the following species:

Table 1: By-catch from pitfall traps Group Genus/Species Invertebrates

Coleoptera (beetles): Silphidae (carrion beetles) Nicrophorus sp.

Staphylinidae (rove beetles) Not assessed to species

Elateridae (click beetles) Hemicrepidius pallidipennis (Mannerheim) H. morio (LeConte) Ctenicera sp. Megapenthes caprella (LeConte) Curculionidae (weevils) Nemocestes incomptus (Horn) Rhyncolus brunneus Mannerheim

Byrrhidae (pill beetles) Not assessed to species

Scolytidae Not assessed to species

Buprestidae Not assessed to species

Cerambycidae Xestoleptura crassipes (LeConte)

Pyrochroidae Ischalia vancouverensis Harrington

Leiodidae (small fungus beetles) Catops sp. Catoptrichus frankenhaeuseri (Mannerheim)

Oedemeridae (false blister beetles) Ditylus gracilis LeConte

Lampyridae Ellychnia hatchi Fender

Cerambycidae (long-horned beetles) Not assessed to species

Zopheridae (used to be Phellopsis obcordata (Kirby) Tenebrionidae)

Tenebionidae (darkling beetles) Ipthiminus serratus Mannerheim Helops pernitens LeConte

109 Throscidae Pactopus hornii (LeConte)

Arachnida: Spiders not assessed to genus or species

Mites not assessed to genus or species

Chilopoda: Millipedes not assessed to genus or species except they were separated into groups: Grey (family Parajulidae), pink (Nearctodesmus insulanus & Keypolydesmus anderisus) and yellow (yellow spotted millipede, Harpaphe haydeniana haydeniana). Centipedes not assessed to genus or species

Hymenoptera: Ants not assessed to genus or species Bees not assessed to genus or species Wasps not assessed to genus or species

Diptera not assessed to genus or species

Heteroptera: not assessed to genus or species

Gastropoda: Slugs not assessed to genus or species Snails not assessed to genus or species

Vertebrates

Mammalia: Shrew Sorex vagrans and Sorex monticolus

Amphibia: Frogs not assessed to genus or species Salamander & Newts not assessed to species

For staphylinids, snails, spiders, crickets, curculionidae, pink millipedes (Nearctodesmus insulanus & Keypolydesmus anderisus) and the millipede Harpaphe haydeniana haydeniana, I made a number of graphs to examine catches across transects

110 within all sites. This allowed me to examine whether there were any large scale differences in abundance a) among old growth and 2nd growth sites, b) by location (e.g. in, out, edge, 5 to 10 m out from edge, 5 to 10 m in from edge) within sites and c) among sites with different patch sizes.

Snails did not show consistent patterns among sites. In old growth sites, they were more common inside of patches, but in 2nd growth sites, they showed generally equal abundance inside patches and in the cut matrices (Figures 1 & 2). No consistent differences by patch size were evident. Snails did show a peak at 5-10 m outside of edges in the 2nd growth sites, a pattern that was seen in some of the beetle species, and which might be related to their becoming trapped in this area due to a generally unwillingness to venture further out into the matrix (Figure 2). Spiders were generally more common in the matrices of both 2nd growth and old growth sites, but did not show consistent patterns with patch size (Figures 3&4). No consistent patterns were apparent for weevils (Figures 5&6). Crickets were similarly abundant in the patches and matrices of 2nd growth sites, but were more abundant in the matrices of old growth sites (Figures 7&8). Pink millipedes were more abundant in the matrices of 2nd growth sites, but were similarly common in the old growth patches and matrices, except for in the large patches of the L VRAM treatment, where they were particularly common (Figures 9&10). Harpaphe millipedes were more abundant in the patches of 2nd growth sites, but were equally abundant among patches and matrices of old growth sites (Figures 11&12). Staphylinid catches were low during 2005, but in both 2nd growth and old growth sites, catches were highest overall in the large patch sites (G1 for 2nd growth, G5 for old growth) and were generally higher in the patches over the matrices (Figures 13&14).

111 Figure 1. Average yearly catches of snails (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Snails

2.0 Into cut matrices Into forest patches G5 1.8 L 1.6 G4 Sm 1.4 M 1.2

1.0

0.8

0.6

Mean no. snails per trap per snails no. Mean 0.4

0.2

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 2. Average yearly catches of snails (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Snails

2.5 Into cut matrices Into forest patches G1 G3L G2 2.0 G3Sm

1.5

1.0

Mean no. snails per trap per snails no. Mean 0.5

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

112

Figure 3. Average yearly catches of spiders (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Spiders

6.0 Into cut matrices Into forest patches G5 L 5.0 G4 Sm 4.0 M

3.0

2.0 Mean no. spiders per trap per spiders no. Mean 1.0

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 4. Average yearly catches of spiders (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Spiders

14.0 Into cut matrices Into forest patches

12.0 G1 G3L 10.0 G2 G3Sm 8.0

6.0

4.0 Mean no. spiders per trap per spiders no. Mean 2.0

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

113 Figure 5. Average yearly catches of cucurlionidae (weevils- all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Curculionidae

2.5 Into cut matrices Into forest patches G5 L 2.0 G4 Sm M 1.5

1.0

Mean no. weevils per trap per weevils no. Mean 0.5

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 6. Average yearly catches of cucurlionidae (weevils- all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Curculionidae

0.9 Into cut matrices Into forest patches 0.8

0.7

0.6 G1 G3L 0.5 G2 0.4 G3Sm

0.3

Mean no. weevils per trap per weevils no. Mean 0.2

0.1

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

114 Figure 7. Average yearly catches of crickets (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Crickets

9.0 Into cut matrices Into forest patches G5 8.0 L G4 7.0 Sm 6.0 M

5.0

4.0

3.0

2.0 Mean no. crickets per trap per crickets no. Mean

1.0

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 8. Average yearly catches of crickets (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Crickets

5.0 Into cut matrices Into forest patches G1 4.5 G3L G2 4.0 G3Sm 3.5

3.0

2.5

2.0

1.5

Mean no. crickets per trap per crickets no. Mean 1.0

0.5

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

115 Figure 9. Average yearly catches of pink millipedes (Nearctodesmus insulanus & Keypolydesmus anderisus) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Pink Millipedes

1.8 Into cut matrices Into forest patches 1.6

1.4 G5 1.2 L G4 1.0 Sm 0.8 M

0.6

0.4 Mean no. millipedes per trap 0.2

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 10. Average yearly catches of pink millipedes (Nearctodesmus insulanus & Keypolydesmus anderisus) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Pink millipedes 0.3 Into cut matrices Into forest patches

0.2 G1 G3L G2 0.2 G3Sm

0.1

0.1 Mean no. millipedes per trap per millipedes no. Mean

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

116 Figure 11. Average yearly catches of Harpaphe haydeniana haydeniana by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Harpaphe haydeniana

0.8 Into cut matrices Into forest patches 0.7

0.6 G5 L 0.5 G4 0.4 Sm M 0.3

0.2 Mean no. millipedes per trap 0.1

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 12. Average yearly catches of Harpaphe haydeniana haydeniana by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Harpaphe haydeniana

1.0 Into cut matrices Into forest patches G1 0.9 G3L G2 0.8 G3Sm 0.7

0.6

0.5

0.4

0.3

0.2 Mean no. millipedes per trap per millipedes no. Mean

0.1

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

117 Figure 13. Average yearly catches of staphylinids (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within old growth group VR sites.

Old Growth Sites Staphylinids 0.5 Into cut matrices Into forest patches 0.5

0.4

0.4 G5 L 0.3 G4 0.3 Sm M 0.2

0.2

0.1 Mean no. staphylinids per trap 0.1

0.0 -30 -25 -20 -15 -10 -5 0 5 10 15 20 25 30 Metres from forest edge

Figure 14. Average yearly catches of staphylinids (all species pooled) by location (In, Out, Edge, 5to 10m in from edge, 5 to 10m out from edge) within 2nd growth group VR sites.

2nd Growth Sites Staphylinids 0.5 Into cut matrices Into forest patches G1 0.4 G3L G2 0.4 G3Sm 0.3

0.3

0.2

0.2

0.1 Mean no. staphylinids per trap 0.1

0.0 -30-25-20-15-10-50 5 1015202530 Metres from forest edge

118

Abundance and Diversity of Millipedes in Managed Forests on Vancouver Island

Tasheena Gardner (200100634) BISC 498 Research Report Department of Biological Sciences Simon Fraser University

December 13, 2005

Supervisor: Dr. Alton Harestad

119 Introduction An adaptive forest management plan introduced by Weyerhaeuser Company Ltd. was designed and implemented to test and demonstrate the effectiveness of variable retention harvesting systems. Part of the management goals was to maintain a functioning ecosystem similar to that in older coastal forests, including its biodiversity.

As well, part of the goals was to identify indicators to assess whether this management approach is effective. The diversity and biomass of invertebrates demonstrates a functioning ecosystem (Black et al. 2001). The sheer numbers, limited mobility and moist habitat preferences of millipedes (i.e., Class Diplopoda) may reveal the state of the ecosystem and therefore be an appropriate ecological indicator of forest conditions. At least 1400 millipede species occur in North America (Shelley 1999). If the abundance and species of millipedes vary with habitat, then millipedes could be used to assess the effects of forest practices on the environment.

The geography of Canada suggests that diverse populations of millipedes occur primarily in British Columbia and Ontario (Shelley 1988). Extensive studies of millipedes in these locations are deficient. A variety of millipede species are distributed along the southern portion of the west coast of British Columbia, a region that includes the study area. I obtained millipede samples consisting of two orders, three families and six species from specific sites on Vancouver Island. Millipedes are scavengers that feed mainly on decaying organic debris on the forest floor and require moist environments to survive. The production of eggs and sperm in millipedes occurs in organs called gonopods, which are associated with the second pair of legs on the millipede (Shelley

1990; Figure 1). The male gonopods are an important characteristic that can be used to

120 differentiate some species found on Vancouver Island because the juveniles of Harpaphe haydeniana look like the adult forms of Tubaphe levii.

I will examine the effects of patch size, edges and forest harvesting on millipede communities by assessing millipede diversity and determining the role that variable retention harvesting systems play in maintaining this diversity. The purpose of this study is:

1) to determine if transect position within a particular forest patch affects the

abundance and species richness of millipedes; and

2) to examine if millipede diversity changes along these gradients on a monthly

basis.

Materials and Methods Millipedes were sampled using pitfall traps because this method is cost-effective and efficient. Pitfall trapping involves placing traps (isopropyl alcohol in cups with a small opening in the lid) along a transect line and collecting the contents of the trap at a later date. Contents of all traps were collected and pooled into the corresponding group location from which they were collected. A total of 201 vials of specimens were collected over a five-month period (May 2005 through September 2005). The pitfall traps were set along transects in different sized variable retention patches radiating from a centre point in the patch to the patch edges, and then up to 25 m beyond the forest edge into the cut matrix (i.e., the clearcut area around a forest patch). These transect positions are identified as: centre, in forest, edge, 5 m out and 25 m out). Depending on the site, four to five pitfall traps were positioned along these transect lines. This design was used in different sized patches in three treatments of second growth forest: G1 (large patches),

121 G2 (small patches) and G3 (2 large patches and 3 small patches). A second growth control group was also sampled. As well, old growth treatments G4 (three small patches) and G5 (2 large patches) contained a similar transect arrangement. Experimental variable retention sites within old growth forests were established as old growth control, old growth patches (small, medium, large) and clearcut sites with a different transect arrangement. Three transect lines were positioned in an equilateral triangular shape so that the placement of the last trap was close to the position where the first trap was set.

Five pitfall traps were set along these triangular transect lines.

Millipedes were collected as by-catch in a study of beetles, so they were not of initial interest when setting the traps. These millipedes were later identified and classified to species. Table 2 lists the species names and their corresponding abbreviations that I used in my report. Specimens were identified by using detailed descriptions of each species provided by Pearsall (2005). When it was difficult to distinguish between juvenile Harpaphe haydeniana and adult forms of Tubaphe levii, I used a dissecting microscope to determine whether or not gonopods were present (Figure 1). If gonopods were present, I classified these millipedes as T. levii, otherwise they were classified as H. haydeniana.

Data relating to the species captured were compiled in Excel spreadsheets and later imported into the Statistical Package for the Social Sciences computer program

(SPSS), which I used for my analyses. The original recorded field transect position for each sample was recoded to make each transect position at a site spatially equivalent among sites. This manipulation permitted a more accurate comparison of the groups. In other words, the centers, edges and positions outside of a forested patch could be

122 compared throughout all sites. Hence, some transects did not have a position at 10 m from the edge (position 2).

The number of millipedes caught each month were divided by the number of traps to account for different trapping effort. Traps were occasionally destroyed by bears or could not be found. In general, the center position at some sites had more traps so they would appear to have a greater total number of millipedes. I had to adjust for this disparate effort because some of this greater abundance was simply due to the greater number of trap samples. The abundance of millipedes was analyzed using a broad scale approach in which I compared the number of individuals for each species relative to the transect position and month in which they were collected. More detailed analyses could be done, but I was limited by my understanding of the study design.

Results A total of 1870 millipedes were collected in this study sample and were comprised of 6 species. I analyzed the relative abundance of different millipede species by their monthly occurrence and their transect position. I conducted a 2-factor ANOVA to determine the significance of the transect position and collection month (Table 1).

Transect position and month are significant factors that influence millipede abundance.

There was no significant statistical interaction between month and transect position.

Hence, I proceeded further with my analyses and examined the abundance of millipedes along transects relative to the collection months.

I created graphs using the information that I identified as being significant.

Figure 2 represents the relative abundance by species at each transect position for all sites combined. The relative composition of the millipede community was similar across

123 transect positions. As well, the same 3 species dominated the community regardless of transect position. Figure 3 better illustrates this dominance of Harpaphe haydeniana,

Tubaphe levii and Keypolydesmus anderisus. The remaining three species, Litiulus alaskanus, Nearctodesmus insulanus and the Parajulids, are relatively minor in terms of relative abundance. I combined all three species to calculate total number of millipedes.

The total number of millipedes was greatest at the center position in a forested patch. The remaining four transect positions had lower numbers of individuals and were roughly the same (Figure 4); however, the standard errors (SE) for these data are large.

I examined the three control groups (clearcut, second growth and old growth) to provide a context for the transect position data and to determine the species that are normally present in these habitat types (Figure 5). Clearcut areas had the lowest numbers of millipedes by species, with Tubaphe levii being the most abundant. Neither the second growth forests nor clearcut areas had Litiulus alaskanus and Nearctodesmus insulanus present whereas Keypolydesmus anderisus and the Parajulids occurred in similar numbers in both locations. Old growth stands had the highest number of species, although Litiulus alaskanus and Nearctodesmus insulanus were rare in these stands. I examined monthly numbers of each species (Figure 6). Litiulus alaskanus was not present until June and remained low in July but increased greatly by August. Conversely, the prevalence of other species tended to decrease after July and then remain low or increase slightly into

September. The numbers of Harpaphe haydeniana are most abundant from May through

July. Their numbers drop substantially after this date and were the least abundant species in September.

124 Discussion Interpretation of results

The objective of my study was to determine the abundance of millipede species by month, site, and transect position at which they were caught. Arthropods are one of the most diverse and adaptive taxa worldwide and represent the vast majority of recognized species in terrestrial ecosystems. Their capacity for rapid response to environmental change allows many arthropods to be useful indicators of change

(Schowalter and Sabin 1991). For this reason, millipedes may be an appropriate indicator species for use in forest management. Different habitat types, such as those where the millipedes were collected, ranged from centres of forest patches and edges of these patches to areas 25 m into clearcuts. The transects were grouped into large or small patch sizes for different sites. My findings indicate that month and relative transect position were important factors that affected millipede abundance, but were independent of each other. Because these factors were independent, their individual effects were worth investigating further. I thought that the month, relative transect position, and overall species composition within these sites was the most pertinent data to analyze in this study because they provide a broad overview about the specimens collected.

Each of the six species identified were present at all transect positions, however, there was a large variation in the relative abundance of each species. There were three dominant species: Harpaphe haydeniana, Tubaphe levii and Keypolydesmus anderisus.

This indicates that the three species were roughly equal among the study sites and suggests that habitat features at the stand scale have little effect on their abundance.

Interestingly, a similar pattern occurs for the remaining three species (Litiulus alaskanus,

125 Nearctodesmus insulanus and the Parajulids), albeit they are less abundant. Perhaps when the three dominant species are present, the three less dominant species are unable to take full advantage of the habitat because the dominant species competitively exclude them. Alternatively, perhaps the three less common species require specific niches that provide them with their survival requirements (i.e., food or microclimate) and these habitat conditions are relatively uncommon within the study sites.

Based upon my results when I compared the total number of individuals (i.e., ignoring species) at each transect point, the center position appeared to have the greatest number of individuals. A plausible explanation for this finding may be the small size of millipedes and that they may respond more to small scale processes than to large scale processes. However, it appears that larger scale effects may be expressed because millipedes were most abundant at the interior-most location but declined near (i.e., 10 m from) the edge and remained low beyond that location. This distance from the edge into a patch could be influenced by microclimate and is well within the zone of an edge effect.

Whatever the habitat changes that are imposed by forest disturbance, they seem to be expressed equally from within a patch and 10 m from the edge into the clearcut. The large confidence intervals occurring at each transect position also has to be considered when interpreting these data because if millipede numbers fluctuate at each site, then the central location may not be as different as it appears when data were combined.

Comparison of the control groups illustrates that there were more millipedes in old growth forests compared to second growth forests, which confirms my expectation.

Old growth forests tend to have a higher diversity of fauna and flora than other forest types due to their habitat complexity. An interesting finding however, was that the

126 species Litiulus alaskanus and Nearctodesmus insulanus were rare in the old growth forest but did not occur in second growth or clearcut. Although these species could be indicators of old forest conditions on Vancouver Island, they are rare and may require too much effort to sample effectively.

The monthly variation of millipede species at a particular site indicates that population numbers may be seasonally dependent. The largest differences I found were for Litiulus alaskanus. This species did not occur until July and then the population increased substantially through to September while the other five species showed an opposite trend. These decreases in the 5 species could be due to seasonal variations in temperature and moisture. Yi and Moldenke (2005) indicated that millipedes were positively associated with litter moisture and that millipedes are indicators of environmental change, in particular with forest thinning. A possible explanation for this seasonal difference among species could be that Litiulus alaskanus has a greater ability to thrive in the drier, hotter months of July and August, thus it increases in prevalence as the other species decline. Because the other 5 species have a similar pattern of decreasing during the hot summer months, they may be more sensitive to moisture gradients and therefore decrease in abundance or avoid surface movements in summer. Pearsall (2005) indicated that millipedes become more active in winter while they seek winter habitats.

This can explain the slight trend for the millipedes to begin increasing in abundance again in September. September is generally cooler and moister and thus greater surface activity could be promoted across all habitat types

Study Design

127 The study design should be reassessed and potentially revised. Initially, the study design appeared to be adequate, however, when I attempted to interpret the data statistically, it became a challenge to accurately express the results in a spatially equivalent manner. For example, Transect Position 2 (10 meters into the forest patch from the edge of the forest) had a lower catch in part because there were fewer traps at this position. Not all treatment groups had five transect points, but instead some had four.

I had to make changes to the data set so each transect point was spatially comparable among sites. Sites with five transect positions had more effort in position 2 than did those with four transect positions because site 2 existed only in a 5 position transect.

There were also discrepancies in design among sites. All groups were set up similarly except for group 3 (G3). G3 had different sized patches within the group, unlike any other group in this study. This made the data more difficult to compare because the data had to be pooled as well as made spatially equivalent. Overall, the study design allowed for the analysis of a large number of millipedes at a broad level, but a more consistent study design would allow for straightforward data interpretation especially at the more specific level.

This research was conducted during the summer months on Vancouver Island.

For a more accurate representation of how changes in forest habitat affect millipede abundance and distribution along a transect, it may be important to incorporate more seasonal variation into the study by collecting data over a longer period. Trends in species richness may be more apparent and affected by more variables if the time span for specimen collection were increased. Moisture or litter composition at each site may also be useful variables to include as co-factors because millipedes rely on moist conditions

128 for their survival and activity. The scope of this study was limited in order to fit within the bounds for a BISC 498 research project. Hence, analysis of these additional variables would have been time consuming and would have increased the compilation time substantially. Patch size was recorded in this study but due to the experimental arrangement and my understanding of this arrangement, I did not include it as a variable in my analyses.

Management Implications

Based upon my findings, millipedes do not appear to be a good indicator species to use when determining the stability or health of a forest ecosystem. Millipedes showed no trend in their abundance with transect location (except for central locations in a patch).

This insensitivity means they would be poor indicators of patch edges compared to clearcuts. The trends that did appear related more to seasonal patterns. Species varied in abundance across the transect locations, but exhibited no defining characteristics that could be used to assess the ecological stability of an area. Further research at these sites and more detailed microhabitat assessments might reveal other factors which would be useful to assess forest management practices.

Sensitivity of millipedes to land management options and seasonal rhythms has been described in previous studies, hence millipedes may still be useful as ecological indicators (Rossi and Blanchart 2005). If millipedes are pursued as an indicator, then their sensitivity to microclimate as well as the interactions among different species should be incorporated into the study design. Millipedes as a group are not restricted to old stands or any particular transect position, however, patch size and corridors of continuous

129 forest may be additional features to consider. It may also be insightful to determine whether the particular species captured are generalists or specialists in the environment.

This could help determine whether a particular species of millipedes, rather than millipedes as a whole, require specific environmental attributes that defines their habitat and indicates ecosystem viability.

Based on this study, it would be premature to imply that millipedes are not good indicators of ecosystem stability even though features such as edge effects and patch size may not predict how millipedes respond. Rather than disregarding the data because it does not seem to ‘fit’ what is expected (e.g., expect clearcuts to have the lowest species diversity and edges to influence diversity significantly in some invertebrates), one could presume millipede fauna are unique. Further research including microhabitat characteristics could be pursued before excluding millipedes from consideration as indicator species.

Acknowledgements

I thank Isobel Pearsall for allowing me to participate in her research project. She kindly provided samples and taught me how to identify millipede species. I would also like to thank Alton Harestad who provided the opportunity for me to expand my knowledge in research while providing his expertise in data analysis and organization, which assisted me in completion of complete my final report.

Literature Cited

Black, S.H., M. Shepard and M.M. Allen. 2001. Endangered invertebrates: the case for greater attention to invertebrate conservation. Endangered Species 18: 1-50. Accessed Nov. 27, 2005. http://www.xerces.org/Endangered/endangered_paper.pdf

130 Pearsall, Isobel. Personal communication, October 2005. Rossi, J.P. and E. Blanchart. 2005. Seasonal and land-use variations of soil macrofauna composition in the Western Ghats, southern India. Soil Biology and Biochemistry 37: 1093-1104. Schowalter, T. D., and T. E. Sabin. 1991. Litter microarthropod response to canopy herbivory, season and decomposition in litterbags in a regenerating conifer ecosystem in western Oregon. Soil Biology Fertility 11: 93–96. Shelley, R.M. 1988. The millipedes of eastern Canada (Arthropoda: Diplopoda). Canadian Journal of Zoology 66: 1638-1663. Shelley, R.M. 1990. A new milliped of the genus Metaxycheir from the Pacific Coast of Canada (Polydesmida: Xystodesmidae), with remarks on the tribe Chonaphini and the western Canadian and Alaskan diplopod fauna. Canadian Journal of Zoology 68: 2310-2322. Shelley, R.M. 1999. Centipedes and millipedes with emphasis on North America fauna. Accessed Nov. 27, 2005. http://www.emporia.edu/ksn/v45n3-march1999/KSNOL45-3.htm Yi, H. and A. Moldenke. 2005. Response of ground-dwelling arthropods to different thinning in young Douglas fir forests of western Oregon. Environmental Entomology 34: 1071-1080.

131 Table 1. Results of one-way ANOVA for millipede abundance (millipede catch per trap) for two factors month and relative transect position.

Tests of Between-Subjects Effects

Dependent Variable: total species effort Type III Sum Source of Squares df Mean Square F Sig. Model 284.781(a) 25 11.391 20.261 .000 Month 21.712 4 5.428 9.654 .000 Relative transect 7.806 4 1.952 3.471 .010 Month * relative 3.647 16 .228 .405 .980 transect Error 89.395 159 .562 Total 374.176 184 a R Squared = .761 (Adjusted R Squared = .724)

Table 2. List of millipede species names used in this report.

Order Species Name Abbreviations Polydesmida Harpaphe haydeniana haydeniana harhay Polydesmida Tubaphe levii tublev Polydesmida Nearctodesmus insulanus nearins Polydesmida Keypolydesmus anderisus keyand Julidae Parajulids paraju Julidae Litiulus alaskanus litala

132

Figure 1. Gonopods of Tubaphe levii that were used to distinguish between the adult form of Tubaphe levii and the juvenile Harpaphe haydeniana. Drawing modified from Shelly (1990).

133

Figure 2. Relative proportions of species in each transect position. Scaled to 100% to give a better representation of the prevalence of each species.

134

species harhay tublev nearins keyand paraju litala

Figure 3. Proportion of species in the total catch of millipedes. Three species are fairly equal in prevalence and the remaining three species occur less often.

135

Number of millipedes t

Figure 4. Mean number (SE) of millipedes per trap at each transect position. Other than the centre of a forest patch, the numbers of millipedes were similar across the transect.

136

Figure 5. Mean number of millipedes per trap in clearcut, second growth and old growth. Nearctodesmus insulanus and Litiulus alaskanus did not occur in clearcut and second growth forest traps, hence bars are not visible.

137

Figure 6. Monthly variation of individuals in each species and percentage of millipedes captured for each month. Litiulus alaskanus was not present until the end of June whereas the other five species tended to decline in number from July onwards.

138

139