in a Northern Hardwood Forest - BioOne

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investigated the impact of logging on native bees (Apoidea) in the Adirondack Mountains of. New York State (USA) in an area isolated from the effects of both ...
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 80(4), 2007, pp. 327–338

Impacts of Logging on Midsummer Diversity of Native Bees (Apoidea) in a Northern Hardwood Forest W. L. ROMEY,1 J. S. ASCHER,2 D. A. POWELL,3

AND

M. YANEK4

ABSTRACT: Although there is a worldwide concern about anthropogenic causes of pollinator declines, little is known about how specific land use practices, such as agriculture, urbanization, and logging, influence the diversity and abundance of native bees. We investigated the impact of logging on native bees (Apoidea) in the Adirondack Mountains of New York State (USA) in an area isolated from the effects of both agriculture and introduced honey bees. Specifically, we measured midsummer bee abundance and diversity on matched 5acre plots two and three years after experimental logging cuts to remove 30, 60, and 100% of the trees. We obtained additional data on floral abundance and weather in relation to bee abundance. We found that bee abundance and diversity was highest in areas where the most trees had been removed. Also, we found little difference in diversity between the two study years, despite major differences in weather. The observed diversity patterns were best explained by the observed increase in abundance of flowering plants such as Rubus and Solidago. Despite the overall relationship between logging and bee diversity, the abundance of four species of Lasioglossum were significantly greater in the 60% tree removal plot than the 100% removal clear cut. Overall, our results suggest that a small-scale tree removal within a northern hardwood forest increases diversity and abundance of many bee species in the short term. KEY WORDS: Apoidea, native bees, land use, forest, pollinator, logging

Native bees are critical to the pollination of many wild plants and are an unrecognized reservoir of agricultural pollinators (Kearnes and Inouye, 1997; Ashman et al., 2004). However, recent studies have shown a historical decline in diversity of native bees which may be linked to anthropogenic causes (Biesmeijer et al., 2006; National Research Council, 2006). Human activity may harm bees in a variety of ways including: pesticide use, land disturbance, and the introduction of exotic species (Aizen and Feinsinger, 1994; Kearnes and Inouye,1997; Cane and Tepedino, 2001; Klein et al., 2002; Kremen, et al., 2002; Goulson, 2003; Kremen, et al., 2004; Tylianakis et al., 2005). In contrast, human or insect activity may benefit some bee species by opening up previously shaded areas and increasing floral resources (Gill, 1996; Michener, 2000). For example, Steffan-Dewenter et al., (2002) found that as the ratio of grassland to temperate forest increased, abundance and diversity of bees increased. Similarly, Winfree et al. (2007) found a negative relationship between forest cover and bee abundance/richness. However, controlled studies of the effects of logging on bee diversity in northern regions are absent. The specific effects of logging on bee populations are difficult to predict. Possible negative impacts include disruption and compaction of soil nest sites, alteration of soil moisture, and loss of natural cavities used as bee nest sites. Positive impacts 1 Dr. W.L. Romey, Department of Biology, State University of New York at Potsdam, Potsdam, NY 13676 USA. e-mail: [email protected], phone: 315-267-2292, FAX: 315-267-3170. 2 Division of Invertebrate Zoology, American Museum of Natural History, New York, NY, 10024, USA. 3 81 S. Casingham, Bexley, OH 43209 USA. 4 Department of Entomology, University of Wisconsin, Madison, WI 53706, USA.

Accepted 15 April 2007; Revised 19 June 2007 E 2007 Kansas Entomological Society

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include more herbaceous flowering plants due to increased light penetration (Aizen and Feinsinger, 1994), warmer soil for soil-nesters (Cane, 1991), and more dead wood for wood-nesting bees. Not only is the type of logging action likely to influence bee diversity, but also the area of the remaining opening should, as predicted by the species/area concept (MacArthur and Wilson, 1967). Therefore, it is important to account for the area logged as a covariate in statistical comparisons, or better, to use equally sized logging cuts in a manipulated experiment. In studies of tropical lepidopteran pollinators (Hill, 1999; Lewis, 2001; Hamer et al., 2003) logging did not significantly influence overall diversity of pollinators. Instead, the number of species remained approximately the same, but there was a turnover in species from ones that were shade-adapted to ones that were adapted to open areas. However, it is difficult to generalize from lepidopteran to bee communities because of the major difference in the needs of their larvae (i.e. leaves vs. pollen). In this study, we tested the impact of different logging practices on the diversity of native bees. In a uniform patch of hardwood forest in the Adirondack mountains in northern New York (isolated from agricultural areas and honey bees) four matching 5-acre plots were treated in one of four ways: 1) left alone (control), 2) 30% of the trees removed, 3) 60% of the trees removed and 4) 100% of the trees removed (clear cut). Two and three years after these logging treatments, we compared bee abundance, diversity, and their preference for different colored traps in midsummer. This study is one of the first to compare the impacts of different logging practices on native bee diversity in a northern hardwood forest.

Methods Study Site Our study was conducted in the Adirondack Park at the FERDA (Forest Ecosystem Research and Demonstration Area), located near the town of Saranac Lake in Franklin County, New York (44u 269 lat and 74u 159 long, 540 m elevation). FERDA is a long-term research and silviculture demonstration area that hosts a variety of ongoing studies (Rechlin et al., 2000). The area is a northern hardwood forest (Smith, 1986) dominated by American beech (Fagus grandifolia Ehrh. (Fagaceae)), sugar maple (Acer saccharum Marsh. (Aceraceae)), and yellow birch (Betula alleghaniensis Britt. (Betulaceae)). Prior to logging, there were no significant pre-existing differences in plant diversity or density among plots (Wade et al., 2003). The 5-acre plots were logged during the winter of 1999–2000 as summarized below (for details see Wade et al., 2003). Plot numbers are listed below (‘‘FERDA plot #’’) for comparison to other published accounts. 1) Control (C): This plot was delineated but not cut. It is heavily shaded (no light reaching forest floor) from June to October. (FERDA plot #3). 2) Single-tree removal (ST): 30% of the tree volume was removed (random tree species). The remaining canopy openings are the size of individual trees. (FERDA plot #4). 3) Shelterwood (SW): 60% of the canopy trees were removed leaving a thin canopy and mostly open, sunny ground. Many dead branches (slash) were left on the ground. (FERDA plot #7).

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4) Clearcut (CC): 100% of trees were removed from the plot leaving no shade. Slash, 15–20 cm in diameter and 0.5 meters in depth, was left covering the ground. (FERDA plot #2). Plots are connected by a narrow, unpaved logging trail, but are otherwise wellseparated from agriculture and urban areas in the center of the Adirondack park buffered by a 50 km radius of woodland and lakes (Jenkins and Keal, 2004). Plots C and ST are adjacent to each other (20 m apart at their closest). Plots SW and CC are also adjacent to each other, but separated from the other two plots by 250 m. Pan traps were used to sample bees (Kirk, 1984; Leong and Thorp, 1999; Giles and Ascher, 2006; Winfree et al., 2007). Similar to these other studies, we used SOLOTM traps (355 ml volume, 15 cm diameter) colored white, yellow, or blue, filled to a depth of 2 cm with a mixture of water and detergent (30 ml Ultra-VistaH dishwashing liquid to 4 liters of dechlorinated tap water). Surveys were conducted approximately every week for five weeks on days that were relatively calm, sunny, and warm, between 19 June 2002–24 July 2002 and 10 June 2003–7 July 2003. In 2002 we started sampling later than 2003 because of a long rainy period. Pan traps were placed every 2 m along two parallel transects 36 m long. Each trap was placed on the highest object within 10 cm of the measured location, on top of slash or rocks whenever possible. These two transects were approximately 30 m apart from each other at the center of 5-acre plots. Each sampling day, 36 pan traps (12 white, 12 yellow, 12 blue) were placed in each of the four plots (144 traps total) at approximately the same time in the morning and left out for six hours (between 10:00–16:00 Eastern Daylight Saving Time during 2002 and 11:00–17:00 during 2003). Traps were placed in approximately the same location every week alternating in a regular pattern (blue, white, yellow). At the end of each afternoon, Apoidea were removed from pan traps, rinsed in 95% ethyl alcohol for 10 minutes, pinned, blown dry, and labeled. On each sampling day, all four plots were sampled, with the start and finish time of each plot within an hour of each other. Traps were collected in the same order they were placed to equalize each trap’s collection time. Over the two years there was a total sampling effort of 8,640 trap-hours: 6 hours 3 36 traps 3 4 plots 3 5 days 3 2 summers. Temperature, rainfall and wind velocity were obtained each sampling day from a nearby (1 km) automated weather station at Paul Smiths College (http://paulsmiths.edu/aai/weather). Flowering plants were surveyed before the logging treatment and monthly every summer thereafter as part of a separate study (Wade et al., 2003). After a thorough search of the plot by botanists, a qualitative abundance score from 0–5 was assigned for each plant (0 5 absent, 1 5 rare, 2 5 infrequent, 3 5 occasional, 4 5 frequent, 5 5 abundant). We obtained a subset of the data from this ongoing work (M. Twery, pers. com., USDA) and analyzed these abundance scores in the following way. We made a list of plant species which were relevant food sources for bees and had received an abundance score of 3 or more on at least one of the floral surveys. Then, the average of the abundance scores for the 30 June 2003 and 29 July 2003 samples was calculated for each plot (Table 1). We then determined, by inspection of the table, whether there was an increasing (+) or decreasing (2) trend in that plant species with increased tree removal (C R CC). The plant species listed here were not necessarily flowering on the dates of the bee surveys.

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Table 1. Three-way ANOVAs on the abundance of bees in 2002 and 2003. Number of bees were transformed as log (x + 1). Data were pooled differently between years leading to different degrees of freedom (see methods). 2002

Plot (P) Week (W) Color (C ) P3W P3C W3C P3W3C Error

2003

ss

df

F

P

ss

df

F

P

18.21 2.35 4.20 4.26 2.28 1.12 1.84 10.67

2 4 2 8 4 8 16 45

38.42 2.47 8.85 2.25 2.40 0.59 0.49

0.000 0.057 0.001 0.041 0.064 0.780 0.941

26.88 13.81 3.33 6.78 1.11 4.48 5.71 51.86

2 4 2 8 4 8 16 225

58.31 14.98 7.23 3.67 1.21 2.43 1.55

0.000 0.000 0.001 0.000 0.309 0.015 0.084

Bees were initially sorted to genus using Rozen (2001) and Michener et al., (1994). Detailed species identifications were made using keys in Mitchell (1960; 1962) and subsequent revisions (listed in Michener, 2000). Difficult species were compared directly to definitively determined specimens in collections, including those of Cornell University and the American Museum of Natural History. Sam Droege (USGS) identified many Lasioglossum (Dialictus) species. Representative voucher specimens of selected species have been deposited in the collections of Cornell University, The American Museum of Natural History, and the USDA, Maryland. Remaining specimens have been retained at the SUNY Potsdam collection. Most specimens were identified to the species level (419 of 526 caught), but for the others, the most precise level of identification was used depending on the level of analysis. Analysis For each year the species diversity was calculated using the Shannon-Weiner Diversity Index (natural log) (Pielou, 1975). In this paper species richness refers to the number of species, and abundance to the number of individuals. At the species level, significance between plots was determined, for those with N . 12, using a x2 test. To test if there was a significant difference in diversity and richness between plots, EcoSim software (Gotelli and Enstminger, 2001) was used to calculate 95% confidence intervals using 1000 independent random resamples of the original data. Additionally, a three-way ANOVA was calculated on the effects of Plot, Week, and Trap Color on the overall abundance of bees. For this analysis we did not directly compare the two years because they were processed differently in the field. Despite equal sampling effort (trap hours) between years, in 2002 the 12 replicate traps for a particular color in one site were pooled into two samples per plot (one for each transect) whereas in 2003 every two traps of a particular color were pooled (six samples per plot). Therefore the 2002 abundance data represent a more conservative method of analysis than 2003. Results The level of tree removal influenced abundance, richness, and diversity (H9) of native bees sampled. Tree removal influenced bee abundance positively (Fig. 1, Table 1). For example, in the unlogged control plot (C), only one individual of one

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Fig. 1. Abundance of the five bee families with respect to treatment plot and year. See methods for plot descriptions.

species (Lasioglossum versans) was trapped over the two summers. Bee abundance was ten times higher in SW than in ST (combined totals for both years: 26 vs. 245 bees, x2 test, P , 0.05). Bee abundance was similar in the SW and CC plots (245 in SW vs. 255 in CC, x2 test, P . 0.05). There were also significant differences in family abundance between plots (Fig. 1, Appendix 1, x2 test, P , 0.05 for each of the five families). There were significant differences between plots for the following species (Appendix 1, where N.12): the stem nesters Ceratina dupla, Hylaeus modestus; the ground nesters Lasioglossum athabascense, L. rufitarse, L. sp.; rotten wood nesters L. cressonii, L. oblongum; and the parasitic bee Sphecodes cressonii. For most of these species comparisons, the lowest abundance (apart from C) was in the ST plot, and the highest in the CC plot. Similarly, there were more Bombus at CC than at other plots (pooled Bombus data, x2 test, P , 0.001). However, Lasioglossum differed from this general trend; it was most abundant in the SW plot. Results are similar in regards to species richness. As tree removal increased, richness increased, from one species in plot C to an average of 36 species in plot CC (Fig. 2). There was significantly greater species richness in CC than in other plots (Fig. 2). There were no significant differences in richness between years at any particular plot (e.g. CC 2002 vs. CC 2003, Fig. 2). In total, 48 species of Apoidea in 12 genera (5 families) were identified (Appendix 1). No honey bees (Apis melifera L.) were caught or seen on any plots during the two summers, suggesting that our community measurements are independent of their effects.

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Fig. 2. Bar chart of species richness and diversity of bees with respect to increased levels of logging (black bars are for 2002, gray bars for 2003). Plot abbreviations are listed in the methods. Bars with a common letter over them are not significantly different; otherwise bars are significantly different at the P , 0.05 level (their 95% confidence intervals calculated by resampling (N 5 1000) do not overlap).

Species diversity (H9) also increased significantly with increased tree removal (Fig. 2). Diversity was significantly higher in CC than in other plots (Fig. 2). Diversity of bees in SW was significantly higher than ST in 2002, but not in 2003. Bee diversity in CC in 2002 was significantly higher than in 2003. However, all other between-plot comparisons of diversity between years were not significant. The logging treatment influenced the floral resources available to bees (Table 2). With an increase in tree removal (C to CC) there was a decrease in eight plant species, many of these spring-flowering herbs (Table 2). However, there was an increase in abundance of five plant species (Table 2), three of which were woody vines, and one a tree (Prunus pensylvanica L.f. (Rosaceae)). Rubus (Rosaceae) was

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Table 2. Mean abundance (5 5 most abundant) of summer flowering plants (arranged by Family) observed during June and July surveys in 2003. The plots are organized from most forested to least (e.g.: C 5 uncut control, ST 5 single-tree removal, SW 5 shelterwood, CC 5 clear cut). Key to abundance values are in methods. Mean Abundance at Plot Flowering Plant Species

Aralia hispida Vent. Aralia nudicaulis L. Solidago spp. L. Viburnum alnifolium Marsh. Clintonia borealis (Ait.) Raf. Maianthemum canadense Desf. Medeola virginiana L. Trillium undulatum Willd. Oxalis acetosella L. Polygonum cilinode Michx. Trientalis borealis Raf. Coptis trifolia (L.) Salisb. Prunus pensylvanica L. f. Prunus serotina Ehrh. Rubus allegheniensis Porter Rubus hispidus L. Rubus idaeus L. Mitchella repens L. Tiarella cordifolia L. Viola sororia Willd. Plant Richness Mean Abundance

Family

C

ST

SW

CC

Trend*

Araliaceae Araliaceae Asteraceae Caprifoliaceae Liliaceae Liliaceae Liliaceae Liliaceae Oxalidaceae Polygonaceae Primulaceae Ranunculaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rubiaceae Saxifragaceae Violaceae

0 3 0.5 5 3 4.5 4 2.5 5 2 3 2 0 2 0 0 1 2 2.5 3

0 3 1 5 2.5 4.5 3 2.5 3 3.5 5 3 1 4 2 0.5 3 3 1 3

1 2 2 4 1 3.5 1.5 2 3 5 2.5 3 2.5 1 4 0.5 5 1 0.5 2

2.5 2 3 2 0.5 3 1 1 3 3 1 1.5 3.5 1 3.5 2.5 5 0.5 1.5 0.5

+ 2 + 2 2 2 2 2 2 0 0 0 + 0 0 + + 0 0 2

16 2.25

19 2.68

20 2.35

20 2.08

* the symbols ‘‘2,o,+’’ represent decreasing, neutral, or increasing flower abundance as amount of tree removal increased.

the dominant flowering plant observed in CC and SW. For example, there were no flowers observed in C during these midsummer collections. SW had the highest abundance of: Coptis trifolia (L.) Salisb. (Ranunculaceae), Polygonum cilinode Michx. (Polygonaceae), and Rubus allegheniensis Porter (Rosaceae) (Table 2). The two summers were quite different in weather and bee abundance, but the richness and diversity of bees did not change substantially between years. In the second year of the bee survey, 36% more bees were collected (322 in 2003 vs. 205 bees in 2002). Despite this, most measures of community structure were very similar between years (Fig. 2). It was cooler and rainier in 2002 than 2003. Specifically, the mean (for the specific sampling days) of the daily maximum (and daily mean) temperature were 22.7uC (16.7uC) in 2002 and 23.6uC (17.8uC) in 2003. Also, there was more than twice as much rain (37.6 cm vs. 17.0 cm) in the three months leading up to our sampling times (April, May and June) in 2002 as in 2003. There was a significant effect of trap color on abundance of bees caught (Table 1). More bees came to white traps than blue or yellow (in order: 252, 163, 112; x2 test, P , 0.001). This appears to be largely due to abundant halictine bees such as Lasioglossum cressonii and L. oblongum. For other species the following significant (x2 test, P , 0.05) color preferences existed: Andrena nivalis (Blue), Ceratina dupla (blue + white . yellow), Sphecodes cressonii (white).

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Discussion Logging led to a significant increase in species diversity, richness, and abundance of native bees sampled in mid-summer in a northern hardwood forest. Improvement in food and nesting resources for the bees are possible causes for this. More Rubus were found in CC and SW plots than in ST and C plots (Fig. 2). The flower of this plant is white, which was also the trap color which attracted the majority of bees in our study. Evidently, removal of trees by logging allowed more sun to reach the forest floor, stimulating growth of flowering herbaceous and weedy plants. Bee abundance is often directly correlated with flower abundance (MacKay and Knerer 1979; Ginsberg, 1983). Other possible causes of the observed relationship between bee diversity and logging include an increase in types of nesting resources (dead wood and weeds) and a greater variety of ground temperatures for larval development (Batra, 1990a; Cane, 1991; Batra 1990b). Weekly variation in abundance was significant (Table 1) but differences in richness and diversity between years was not (Fig. 2). Short term differences were probably due to the weather and the emergence of different species of bees. However, it was interesting that there was no significant change in bee richness and diversity in SW and CC plots between 2002 and 2003 (Fig. 2) and no significant change in diversity (H9) in ST and SW plots (Fig. 2). However, there was a significant, but small, decrease in diversity in the CC plot between 2002 and 2003. These small differences in diversity between the two years for each type of plot suggest that the community measures are robust to major changes in weather and that one or two summers of systematic sampling may be enough to obtain reliable estimates of a community (Williams et al., 2001) for comparisons between areas and over years. There are several limitations of our findings which should be taken into mind. First, although pan trapping is a statistically sound way of standardizing sampling effort among plots, it is biased in favor of smaller species (Cane et al., 2000; Giles and Ascher, 2006), underestimating the stronger flying genera such as Bombus, Megachile, and Colletes. However, this bias in sampling would have been consistent across the treatments. Also, the number of species we observed may be lower than in other studies, because the forest was recently cleared; little is known about patterns of bee colonization to such areas. Second, we sampled during the middle of the summer and missed the bees that emerge in the spring and fall. However, the summer is relatively short in the Adirondack Mountains; the average growing season is only about 100 days in many areas (Jenkins and Keal, 2004). Therefore we felt that this mid-summer sample might provide a good community average while acknowledging that it does not reflect the total species richness. At a global scale it appears that many human activities have a negative impact on bee populations across habitat types (as reviewed in Biesmeijer et al., 2006). But for logging, our findings show that removal of trees increased bee diversity. This supports the few other recent studies on this relationship (Steffan-Dewenter et al., 2002; Winfree et al., 2007). For many species of bees, we might consider the maintenance of trees (the other side of forestry), rather than their removal, as a potentially negative anthropogenic influence. It is not clear whether the positive relationship between logging and bee diversity that we found is a short or long-term effect. There may be a successional effect (MacKay and Knerer, 1979) of opportunistic species of bees coming into a recently logged area. Studies on other orders of insects show a rapid rise in species after a disturbance for the first few

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years, then an overall decrease in diversity (Hawkins et al., 1982). A final caveat is that our 5-acre clearings had a relatively high edge/area ratio which provides many niches for bees in the ecotone (Cane, 2001). Commercial logging operations would typically clear larger areas and the decreased edge/area ratio would not lead to as great an increase in bee diversity. Therefore, care should be taken before assuming that logging leads to a sustained increase in bee diversity; long-term studies at different scales are warranted. In conclusion, our study supports the hypothesis that abundance, richness and diversity of native bees increases relative to the number of trees removed from a northern hardwood forest. The overall increase is mirrored by the increase in flowering woody vines/shrubs during the middle of the summer, but may also have to do with increased nesting sites. Other studies on seasonal and yearly fluctuations of this phenomenon in a variety of forest types would be beneficial in understanding the impact of logging on bee populations. Acknowledgements The research was funded by an NSF-REU fellowship through G. Gilchrest, T. Langen, and S. Grimberg at Clarkson University. Additional facilities and support was provided by SUNY Potsdam and a Merwin Rural Services Institute Grant to WLR. We thank S. Droege for help identifying the Halictidae and J. Terhune for statistical advice. We also thank J.C. Biesmeijer, S. Batra, and J. Schreer for improving the manuscript with their comments. For permission to use the study area, we thank the USDA Forest Service and Paul Smith’s College. And within the USDA we thank the following for the floral data: M. Twery for study design and implementation, G. Wade for study design, and J. Rapp for data collection. Neither the authors nor their institutions endorse any proprietary products mentioned in this paper. Literature Cited Aizen, M. A., and P. Feinsinger. 1994. Habitat fragmentation, native insect pollinators, and feral honey bees in Argentine ‘Chaco Serrano.’ Ecological Applications 4:378–392. Ashman, T.-L., T. M. Knight, J. A. Steets, P. Amarasekare, M. Burd, D. R. Campbell, M. R. Dudash, Michele, M. O. Johnston, S. J. Mazer, R. J. Mitchell, J. Randall, M. T. Morgan, and W. G. Wilson. 2004. Pollen limitation of plant reproduction: ecological and evolutionary causes and consequences. Ecology 85:2408–2421. Biesmeijer, J. C., S. P. M. Roberts, M. Reemer, R. Ohlemuller, M. Edwards, T. Peeters, A. P. Schaffers, S. G. Potts, R. Kleukers, C. D. Thomas, J. Settele, and W. E. Kunin. 2006. Parallel declines in pollinators and insect-pollinated plants in Britain and the Netherlands. Science 313:351–354. Batra, S. W. T. 1990a. Bionomics of a vernal solitary bee Andrena (Scrapteropsis) alleghaniensis Viereck in the Adirondacks of New York (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society 63:260–266. Batra, S. W. T. 1990b. Bionomics of Evylaeus comagenensis (Knerer and Atwood) (Halictidae), a facultatively polygynous, univoltine, boreal halictine bee. Proceedings of the Entomological Society of Washington 92:725–731. Cane, J. H. 1991. Soils of ground-nesting bees (Hymenoptera: Apoidea): Texture, moisture, cell depth and climate. Journal of the Kansas Entomological Society 64:406–413. Cane, J. H. 2001. Habitat fragmentation and native bees: a premature verdict? Conservation Ecology 5(1):3. On-Line: http://www.consecol.org/vol5/iss1/art3.(Last accessed 12 April 2007). Cane, J. H., and V. J. Tepedino. 2001. Causes and extent of declines among native North American invertebrate pollinators: detection, evidence, and consequences. Conservation Ecology 5(1), OnLine: http://www.consecol.org/vol5/iss1/art1. (Last accessed 12 April 2007).

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Cane, J. H., R. L. Minckley, and L. J. Kervin. 2000. Sampling bees (Hymenoptera: Apiformes) for pollinator community studies: pitfalls of pan-trapping. Journal of the Kansas Entomological Society 73:225–231. Giles, V., and J. S. Ascher. 2006. A survey of the bees of the Black Rock Forest Preserve, New York (Hymenoptera: Apoidea). Journal of Hymenoptera Research 15:208–231. Gill, D. E. 1996. The natural population ecology of temperate terrestrials: Pink Lady’s Slipper, Cypripedium acaule. In C. Allen (ed.). N.A. native terrestrial orchids, N.A. Native Orchid Conf, pp. 91–100. Ginsberg, H. S. 1983. Foraging ecology of bees in an old field. Ecology 64:165–175. Gotelli, N. J., and G. L. Entsminger. 2001. EcoSim: Null model software for ecology. Version 7.0, Acquired Intelligence Inc. and Kesey-Bear. On-Line: http:/homepages.together.net/˜gentsmin/ ecosim/ecosim.htm. (Last accessed 12 April 2007). Goulson, D. 2003. Effects of introduced bees on native ecosystems. Annual Review of Ecological and Evolutionary Systems 34:1–26. Hawkins, C. P., M. L. Murphy, and N. H. Anderson. 1982. Effects of canopy, substrate composition, and gradient on the structure of macroinvertebrate communities in Cascade Range streams of Oregon. Ecology 63:1840–1856. Hamer, K. C., J. K. Hill, S. Benedick, N. Mustaffa, T. N. Sherratt, M. Maryati, and V. K. Chey. 2003. Ecology of butterflies in natural and selectively logged forests of northern Borneo: the importance of habitat heterogeneity. Journal of Applied Ecology 40:150–162. Hill, J. K. 1999. Butterfly spatial distribution and habitat requirements in a tropical forest: impacts of selective logging. Journal of Applied Ecology 36:564–574. Jenkins, J., and A. Keal. 2004. The Adirondack Atlas. Syracuse University Press, Syracuse, NY. 275 pp. Kearns, C. A., and D. W. Inouye. 1997. Pollinators, flowering plants, and conservation biology. BioScience 47:297–308. Kirk, W. D. J. 1984. Ecologically selective coloured traps. Ecological Entomology 9:35–41. Klein, A. M., I. Steffan-Dewenter, D. Buchori, and T. Tscharntke. 2002. Effects of land-use intensity in tropical agroforestry systems on coffee flower-visiting and trap-nesting bees and wasps. Conservation Biology 16:1003–1014. Kremen, C., N. M. Williams, and R. W. Thorp. 2002. Crop pollination from native bees at risk from agricultural intensification. Proceedings of the National Academy of Sciences 99:16812–16816. Kremen, C., N. M. Williams, R. L. Bugg, J. P. Fay, and R. W. Thorp. 2004. The area requirements of an ecosystem service: crop pollination by native bee communities in California. Ecology Letters 7:1109–1119. Lewis, O. T. 2001. Effect of experimental selective logging on tropical butterflies. Conservation Biology 15:389–400. Leong, J. M., and R. W. Thorp. 1999. Colour-coded sampling: the pan trap colour preferences of oligolectic and nonoligolectic bees associated with a vernal pool plant. Ecological Entomology 24:329–335. MacKay, P. A., and G. Knerer. 1979. Seasonal occurrence and abundance in a community of wild bees from an old field habitat in southern Ontario. Canadian Entomology 111:367–376. MacArthur, R. H., and E. O. Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, N.J. 203 pp. Michener, C. D. 2000. The Bees of the World. Johns Hopkins University Press, Baltimore, Maryland. 913 pp. Michener, C. D., R. J. McGinley, and B. N. Danforth. 1994. The Bee Genera of North and Central America. Smithsonian Institution Press, Washington, D.C. 209 pp. Mitchell, T. B. 1960. Bees of the Eastern United States Volume 1, North Carolina Agricultural Experiment Station, NC, Technical Bulletin, 141. Mitchell, T. B. 1962. Bees of the Eastern United States Volume 2, North Carolina Agricultural Experiment Station, NC, Technical Bulletin, 152.. National Research Council. 2006. Status of Pollinators in North America. National Academy Press, Washington, D.C. Pielou, E. C. 1975. Ecological Diversity. Wiley, New York. Rechlin, M., M. Twery, G. Wade, and M. Storey. 2000. The other half of a ‘‘forever wild’’ park: Education and research at the Adirondack Visitor Interpretive Center. Adirondack Journal of Environmental Studies Fall:5–10.

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Rozen, J. G. 2001. Syllabus of the bee genera of the New England and Mid-Atlantic Regions, extracted from Michener, C. D., J. McGinley, and B. N. Danforth. 1994. The bee genera of North and Central America. Smithsonian University Press, Washington, D.C. 209 pp. Smith, D. M. 1986. The Practice of Silviculture. John Wiley and Sons, Inc., 527 pp. Steffan-Dewenter, I., U. Munzenberg, C. Burger, C. Thies, and T. Tscharntke. 2002. Scale-dependent effects of landscape context on three pollinator guilds. Ecology 83:1421–1432. Tylianakis, J. M., A. M. Klein, and T. Tscharntke. 2005. Spatiotemporal variation in the diversity of hymenoptera across a tropical habitat gradient. Ecology 86:3296–3302. Wade, G. L., J. A. Myers, C. R. Martin, K. Detmar, W. Mator, M. J. Twery, and M. Rechlin. 2003. Vascular plant species of the Forest Ecology Research and Demonstration Area, Paul Smith’s, New York. U.S.D.A. Northeastern Research Station Note NE, publ. #380. Winfree, R., T. Griswold, and C. Kremen. 2007. Effect of human disturbance on bee communities in a forested ecosystem. Conservation Biology 21:213–223. Appendix 1. Abundance of bee species at four different types of logging plots in 2002 and 2003. C is the control, ST 5 single-tree cut, SW 5 shelterwood, CC 5 clear cut. N is the total number of species caught in both years. P is the chi-square significance level comparing the combined (year 2002 + 2003) three noncontrol plots. P was only calculated for species where N . 12. (* , 0.05, ** , 0.01, *** , 0.001). 2002 Family

Genus Species

Andrenidae Andrena crataegi Robertson A. geranii Robertson A. nigrihirta (Ashmead) A. nivalis Smith A. regularis Malloch A. rufosignata Cockerell A. thaspii Graenicher A. wheeleri Graenicher A. wilkella (Kirby) (Family) Apidae Anthophora terminalis Cresson Bombus rufocinctus Cresson B. ternarius Say B. vagans Smith Ceratina dupla Say Nomada cuneata (Robertson) N. pygmaea Cresson (Family) Colletidae Hylaeus basalis (Smith) H. annulatus (Linnaeus) H. modestus Say (Family) Halictidae Augochlorella aurata (Smith) Halictus ligatus (Say) Lasioglossum acuminatum (McGinley) L. athabascense (Sandhouse) L. cattellae (Ellis) L. coriaceum (Smith) L. cressonii (Robertson) L. divergens (Lovell) L. dreisbachi (Mitchell) L. foxii (Robertson) L. heterognathum (Mitchell) L. leucozonium (Schrank)

2003

c

st

sw

cc

c

st

sw

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0

0 0 1 7 0 1 0 0 0 9 0 1 0 1 1 0 0 3 0 0 2 2 1 0 0

1 0 1 6 2 3 2 1 3 19 2 0 3 4 1 0 0 10 2 1 3 6 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 1 2 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 4 3 0 0 0 1 0 8 0 0 0 1 1 0 0 2 0 0 2 2 0 1 0

0 0 0 0 0 0 0 0 0

0 0 0 2 0 0 0 0 0

1 3 0 24 4 0 0 0 0

6 2 0 7 4 0 1 1 1

0 0 0 0 0 0 0 0 0

0 0 0 1 0 0 0 0 0

1 6 4 46 3 0 0 0 2

cc

N

0 1 2 4 0 0 0 4 0 11 0 0 2 3 10 2 1 18 5 1 6 12 2 3 1

1 1 10 22 2 4 2 6 3

P

*

*** 2 1 5 9 13 2 1

**

*** 7 2 14

* ***

4 4 1

9 17 1 12 7 11 32 112 1 12 1 1 1 2 0 1 0 3

***

***

338

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY Appendix 1. Continued. 2002

Family

Genus Species

L. nigroviride (Graenicher) L. oblongum (Lovell) L. obscurum (Robertson) L. rufitarse (Zetterstedt) L. (Dialictus)# L. versans (Lovell) L. viridatum (Lovell) Sphecodes banksii Lovell S. clematidis Robertson S. coronus Mitchell S. cressonii (Robertson) S. sp. S. mondibularis Cresson (Family) Megachilidae Hoplitis producta (Cresson) H. spoliata (Provancher) Megachile gemula Cresson M. melanophaea Smith M. relativa Cresson Osmia albiventris Cresson O. felti Cockerell O. proxima Cresson O. pumila Cresson O. sp. (Family) Grand Totals

2003

c

st

sw

cc

c

st

sw

cc

N

0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1

0 9 0 0 2 0 0 0 0 0 0 0 0 13 0 0 0 0 0 0 0 0 0 0 0 15

0 23 1 10 8 1 1 0 0 0 3 0 0 80 0 0 0 0 0 0 0 0 0 0 0 94

0 15 0 3 5 6 2 0 1 1 2 0 0 58 0 0 1 0 1 0 0 0 0 0 2 95

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 1 0 1 6 39 20 112 0 0 0 1 1 4 4 22 0 7 4 26 0 0 0 7 0 0 4 7 0 4 1 5 0 0 0 1 0 0 0 1 0 5 6 16 0 3 3 6 0 4 2 6 8 130 102 0 1 3 4 0 0 1 1 0 1 1 3 0 0 1 1 0 0 0 1 0 1 1 2 0 0 1 1 0 5 7 12 0 1 1 2 0 0 1 1 0 9 17 11 151 160 527

P

*** ** **

*

***

*** ***