Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 379 2019 NORTHEASTERN NATURALIST 26(2):379–391 Wild Bees (Hymenoptera: Apoidea) of the Ossipee Pine Barrens Erika M. Tucker1 and Sandra M. Rehan1,* Abstract - The pine barrens of the American Northeast is a globally rare and endangered landscape. The handful of pine barrens remaining in the Northeast provide valuable habitat to many threatened and endangered plant and animal (including insect) species, yet little is known about the bee fauna in this habitat. Here we present the results of the first faunal bee survey of the Ossipee Pine Barrens (OPB) in New Hampshire. We collected over 800 specimens from pan and sweep samples representing 95 species. We documented 1 species, Megachile mucida, for the first time in the state of New Hampshire and recorded 4 introduced species. In addition to general surveying, we surveyed landscapes with 4 different types of management for the OPB. Landscapes that incorporated both burning and mowing after burning into the management regime supported bee communites that had significantly greater abundance than all other treatments and greater species richness than all other landscapes, though not signficantly higher than those that were mowe d without burning. Introduction Northeastern Pinus (pine) barrens are characterized by nutrient-poor, fire-dependent forest ecosystems dominated by Pinus rigida Mill. (Pitch Pine; Pinaceae) and Quercus ilicifolia Wangenh. (Scrub Oak; Fagaceae). These ecosystems are globally rare (G2-ranked community type: 6–20 occurrences worldwide) and only represented by a handful of forests scattered across the Northeast (Maine, New Hampshire, Massachusetts, New York, and Pennsylvania) (Bried and Dillon 2012, Bried et al. 2014, Howard et al. 2011, Patterson et al. 2016). Pine barrens are known to support habitat specialists and help sustain some rare and endangered plants and animals such as Antrostomus vociferus Wilson (Eastern Whip-poor-will), Sylvilagus obscurus Chapman, Cramer, Dippenaar, & Robinson (Appalachian Cottontail), Plebejus melissa samuelis Nabokov (Karner Blue Butterfly), and Callophrys irus (Godart) (Frosted Elfin Butterfly) (Bried et al. 2014, Leuenberger et al. 2016, NatureServe 2017, Swengel 1998). Fire-dependent habitats, such as pine barrens, also have the potential to provide bee-friendly habitat (Bried and Dillon 2012). Forests and grasslands that are thinned periodically by burning provide more open ground for nesting sites, can stimulate growth of annual flowers and prolong flowering periods (Campbell et al. 2018, Grundel et al. 2010, Mola et al. 2018, Wrobleski and Kauffman 2003). In burned pine forest habitats in Greece, Turkey, and Finland, bee abundance was found to be significantly greater than in non-burned pine forest habitats (Potts 2006, Rodriguez and Kouki 2015). Similarly, in California grasslands, Bombus 1Department of Biological Sciences, University of New Hampshire, Durham, NH 03824. *Corresponding author - sandra.rehan@unh.edu. Manuscript Editor: David Orwig Northeastern Naturalist 380 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 spp. (bumble bees) and floral abundance were also found to be greater in burned compared to non-burned sites (Mola et al. 2018). While burned pine forests and grasslands are not the same as pine barrens, they give some insight into the effect fire has on bees in fire-dependent ecosystems. The Ossipee Pine Barrens (OPB) of New Hampshire is one of the largest pine barrens in New England and is considered an endangered landscape (Howard et al. 2011, The Nature Conservancy 2018). Consisting of about 1093 ha, it is situated across 5 towns in Carroll County, NH: Effingham, Freedom, Madison, Ossipee, and Tamworth (Howard et al. 2011, The Nature Conservancy 2018). Despite the potential to support rare and unique species, there is still little known about the pine barrens’ wild bee community (Bried and Dillon 2012, Wheeler 1991). Here we document the first faunistic survey of wild bees in the (OPB). The objectives of this study were to: (1) provide a contemporary survey and species checklist of the wild bees currently inhabiting the OPB, (2) evaluate how bee species composition in the OPB varies from that in nearby regions, and (3) determine how wild bees are affected by different habitats and management regimes within the pine barrens. Methods Field-site description We sampled wild bees from 27 sites across 7 towns (Fig. 1, Table 1) in northern New Hampshire in the OPB. These sites were concentrated in and around The Nature Figure 1. Inset: Location of the Ossipee Pine Barrens within New Hampshire. The asterisk marks Strafford County, NH, and the double triangles mark the White Mountains National Forest. The square indicates the location of the Ossipee Pine Barrens. Enlarged section: Map of the treatment areas sampled in the Ossipee Pine Barrens, with A–J corresponding to coordinates listed in Table 1 (treatment map provided by the Nature Conservancy). Burned and then mowed treatments are in blue, burned-only in orange, control in grey, and mowedonly in green. Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 381 Conservancy land (43.78°N, 71.10°W), covering an area of ~5.90 km² with an average elevation of 195 m. We assigned 9 of these sites to a specific treatment based on management regime information and maps provided by The Nature Conservancy (Fig. 1). Treatments and sample size (n) are as follows: burned then mowed (2), burned only (3), mowed only (1), and unmanaged wild areas designated as controls (3). The only control site to have any human management was Site H, which was a gravel pit and is now a restoration area that has been replanted since 2012–2013 with species that normally occur in in the understory of this pine barrens habitat, including Pitch Pine, Scrub Oak, and some Vaccinium (blueberry) and Schizachyrium scoparium (Michx.) Nash (Little Bluestem). It has not been mowed, burned, or otherwise altered except for replanting. All burned sites (including burned-only sites) underwent a single pre-burn mow to control fuel load and make the burn safer. Burned and then mowed sites were mowed again ~1 yr after the burn, with Table 1. Ossipee Pine Barrens collection site information. Abund. = abundance, Rich. = richness, Elev. = elevation. Coordinates Bee Elev. # Location Treatment Nearest town (°N, °W) Abund. Rich. (m) 1 3 Plains Road - Tamworth 43.887, 71.178 2 2 246 2 Bear Camp - Ossipee 43.798, 71.158 15 7 179 3 Burke Field - Tamworth 43.896, 71.155 95 19 246 4 Camp area pasture - Tamworth 43.846, 71.257 57 13 246 5 Deer Hill Road - Tamworth 43.887, 71.184 20 11 246 6 Eaton swimming pond - Eaton 43.900, 71.044 1 1 285 7 Goodwin town forest - Madison 43.852, 71.153 80 29 187 8 Junction of Highways - Effingham 43.792, 71.035 18 10 151 28 and 153 9 Lily Pond - Madison 43.836, 71.187 26 16 187 10 Madison Lot - Silver Lake 43.866, 71.187 44 24 180 11 Mason Road - Ossipee 43.804, 71.056 18 11 179 12 McDonalds - Tamworth 43.820, 71.205 3 3 246 13 Ossipee Road - Ossipee 43.794, 71.054 5 3 179 14 Ossipee Valley - Ossipee 43.832, 71.258 10 3 179 Cemetery 15 Pizza Barn - Ossipee 43.793, 71.178 12 9 179 16 Site A Burned Freedom 43.822, 71.162 34 21 139 17 Site B Mowed Freedom 43.831, 71.178 18 14 139 18 Site D Burned Madison 43.839, 71.182 5 4 187 19 Site E Burned Freedom 43.833, 71.173 29 13 139 20 Site F Burned Madison 43.837, 71.171 113 27 187 then mowed 21 Site G Burned Tamworth 43.837, 71.173 32 18 246 then mowed 22 Site H Control Freedom 43.831, 71.169 35 21 139 23 Site I Control Freedom 43.828, 71.175 2 4 139 24 Site J Control Madison 43.846, 71.185 3 3 187 25 Smith Flats - Tamworth 43.833, 71.236 119 21 246 26 Stillings Lane - Freedom 43.793, 71.066 16 10 139 27 Unamed road - Ossipee 43.829, 71.173 1 1 179 Northeastern Naturalist 382 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 Site F having been burned in August 2009 and 2013 and Site G in September 2007 and October 2016. Of the burned-only sites, site A was burned in October 2013, while site D and E were burned in September 2010. The mowed-only site, Site B, was mowed in late fall 2016. We estimated site elevations with FreeMapTools (https://www.freemaptools.com/elevation-finder.htm) using the GPS coordinates of collection sites as input. We conducted sampling on 24 June 2017 as part of the New England Bee Bioblitz annual event by 15 people for an estimated 45 working hours of sweep netting. It was an overcast morning and sunny afternoon. We employed about 300 standard pan traps for passive collection of bees. These traps were of alternating colors (white, yellow, blue), filled with soapy water, and set out 10 m apart in a transect line for ~6 h. Transect lines were in short vegetation or cleared areas to allow bees to find the traps. We also actively collected bees using standard Bioquip (Bioquip Pruducts, Inc., Rancho Dominguez, CA) collapsible aerial nets for sweeping. We sampled all possible flowering plant blooms throughout collecting locations. We excluded from analyses all Apis mellifera L. (Western Honey Bee; Apidae) specimens because the focus of the study was on unmanaged wild bees. We labeled with locality information and stored in vials of 70% ethanol specimens from both methods of collection. Curation and preservation We washed, dried, and processed all specimens, to each of which we assigned a unique QR code identification number following procedures in Droege (2015) and Tucker and Rehan (2017a, b). We identified all specimens to species using the following identification guides: Gibbs (2011) for metallic Lasioglossum (Halictidae), Mitchell (1960, 1962) for Nomada, Rehan and Sheffield (2011) for Ceratina (Apidae), and www.DiscoverLife.org (Ascher and Pickering 2017; referenced in DiscoverLife: Coelho 2004, Grigarick 1968, Laverty and Harder 1988, Michener 2000, Michener et al. 1994, Mitchell 1980, Ordway 1966, Plowright and Pallett 1978) for all other genera. To identify any potential new state records, we compared our lists to those of Bartomeus et al. (2013), Tucker and Rehan (2016, 2017a, 2018), www.DiscoverLife.org (Ascher and Pickering 2017), and the digital records available from the University of New Hampshire Insect Collection database (UNHC 2018). We used the Very Handy Bee Manual (Droege 2015) to identify introduced bee species. Voucher specimens and associated data are deposited in the University of New Hampshire Insect Collection (Durham, NH). Analyses We compiled datasets to compare species richness variation among the OPB, the White Mountain National Forest (WMNF; Tucker and Rehan 2017a), and longterm surveys conducted in Strafford County, NH (Tucker and Rehan 2017b, 2018; UNHC 2018). We employed the interactive tool VENNY (Oliveros 2007) to visualize variation in species coexistence and exclusivity in relation to different habitats. We compiled additional datasets to examine abundance and richness variation of bee species for management regimes in the OPB. We used generalized linear Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 383 models (GLM) with bee abundance and bee species as variables under a Poisson distribution and performed statistical analyses in R (R Core Team 2015) with the associated packages ‘ggfortify’ (Horikoshi and Tang 2015) and ‘multcomp’ (Hothorn et al. 2008). We subsequently performed a post-hoc Tukey analysis on any significant results. Results Bees of the OPB Pan trap and sweep net samples collected a total of 815 bee specimens. These specimens represented 5 families, 19 genera, and 95 species (Table 2). Halictidae was the most abundant and diverse family, with 405 specimens representing 33 species. Abundance of Halictidae was largely comprised of Halictus ligatus, with 104 specimens. Andrenidae was the next most abundant family, with 182 specimens representing 19 species, and Megachilidae was the second most diverse, with 22 species represented in the 94 specimens collected. We collected 112 specimens of Apidae representing 16 wild bee species. We collected 5 species and 22 specimens of Colletidae, all in the genus Hylaeus. In the OPB, bees were most abundant in Smith Flats, while we detected the greatest richness in Goodwin Town Forest (Table 1).We recorded the species Megachile mucida for the first time in the state of New Hampshire during the course of this survey. Additionally, we found 3 introduced bee species, Andrena wilkella, Lasioglossum leucozonium, and Anthidium oblongatum (Table 2). OPB bee similarity to nearby regions Despite the relative close proximity of OPB to the WMNF (~48 km), bee species composition varied greatly between the 2 regions. Tucker and Rehan (2017a) sampled bees from the WMNF with similar protocols at about the same time of year. Of the 95 species collected in the OPB, we found 30 species exclusively in the OPB and 65 species in both regions (Fig. 2). Species composition was also notably different than in Strafford County, NH (~97 km south of OPB), where thorough long-term bee surveys have been conducted (Tucker and Rehan 2017b, 2018). In comparison to Strafford County, we found 7 bee species sampled only at the OPB and 88 species present in both Strafford County and the OPB (Fig. 2). In comparison with both the WMNF and Strafford County, 5 species were only found in the OPB (Andrenidae: 1, Megachilidae: 2, Apidae: 2; Fig. 2). There were 2 species of Andrenidae—Andrena brevipalpis and Andrena ceanothi—found in the OPB and WMNF, but not in Strafford County despite extensive surveying in that area (Fig. 2). Management regime effects We collected a total of 271 bees representing 59 species from 9 samples, across 4 treatments (Table 1; Sites A–J). Bee abundance was significantly affected by management regime (χ2 = 129.33, df = 5, P = less than 0.001; Fig. 3). Post-hoc analyses examining management revealed that burned and then mowed treatments had significantly more bees than any others. Control treatments had the lowest bee Northeastern Naturalist 384 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 Table 2. Species checklist for all bees species recorded in the Ossipee Pine Barrens (OPB) during this study. Species previously found (+) in the White Mountain National Forest (WMNF; Tucker and Rehan 2017a) and Strafford County, NH (Tucker and Rehan 2017b, 2018), are also listed. * = introduced species; † = a new species record for the state of NH. [Table continued on following page.] Family/OPB species list Abundance Relative abundance WMNF Strafford Andrenidae Andrena alleghaniensis Viereck 1 0.12% + + Andrena brevipalpis Cockerell 1 0.12% + Andrena carlini Cockerell 17 2.09% + Andrena ceanothi Mitchell 23 2.82% + Andrena cressonii Robertson 1 0.12% + + Andrena dunning Cockerell 5 0.61% + Andrena erigeniae Robertson 1 0.12% + Andrena imitatrix Cresson 2 0.25% + + Andrena krigiana Robertson 9 1.10% + + Andrena melanochroa Cockerell 1 0.12% + Andrena morrisonella Viereck 1 0.12% + Andrena nivalis Smith 1 0.12% + + Andrena robertsonii Dalla Torre 4 0.49% + + Andrena rugose Robertson 1 0.12% + Andrena vicina Smith 7 0.86% + Andrena virginiana Mitchell 1 0.12% Andrena w-scripta Viereck 1 0.12% + + *Andrena wilkella Kirby 95 11.66% + + Calliopsis andreniformis Smith 10 1.23% + + Apidae Bombus bimaculatus Cresson 40 4.91% + + Bombus griseocollis De Greer 13 1.60% + + Bombus impatiens Cresson 5 0.61% + + Bombus perplexus Cresson 3 0.37% + + Bombus sandersoni Franklin 23 2.82% + + Bombus ternarius Say 8 0.98% + + Ceratina calcarata Robertson 3 0.37% + + Ceratina dupla Say 1 0.12% + + Ceratina mikmaqi Rehan & Sheffield 3 0.37% + + Nomada articulata Smith 3 0.37% + + Nomada cf. bethunei Grote & Robinson 2 0.25% Nomada cressonii Cockerell 2 0.25% + Nomada depressa Cresson 2 0.25% + + Nomada illinoensis Robertson 1 0.12% Nomada lepida Cresson 2 0.25% + Nomada maculata Cresson 1 0.12% + Colletidae Hylaeus affinis Smith 1 0.12% + + Hylaeus annulatus L. 3 0.37% + + Hylaeus illinoisensis (Robertson) 1 0.12% + Hylaeus mesillae (Cockerell) 2 0.25% + + Hylaeus modestus Say 15 1.84% + + Halictidae Agapostemon texanus Cresson 14 1.72% + + Agapostemon virescens (Fabricius) 14 1.72% + + Augochlora pura (Say) 1 0.12% + + Augochlorella aurata (Smith) 24 2.94% + + Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 385 Table 2, continued. Family/OPB species list Abundance Relative abundance WMNF Strafford Augochloropsis metallica (Fabricius) 3 0.37% + Halictus confusus Smith 11 1.35% + + Halictus ligatus Say 104 12.76% + + Halictus rubicundus (Christ) 1 0.12% + + Lasioglossum abanci (Crawford) 1 0.12% + + Lasioglossum acuminatum McGinley 15 1.84% + + Lasioglossum admirandum (Sandhouse) 2 0.25% + + Lasioglossum birkmanni (Crawford) 2 0.25% + + Lasioglossum cressonii Robertson 47 5.77% + + Lasioglossum ephialtum Gibbs 29 3.56% + + Lasioglossum imitatum (Smith) 4 0.49% + + Lasioglossum leucocomum (Lovell) 3 0.37% + + *Lasioglossum leucozonium Schrank 2 0.25% + + Lasioglossum nigroviride (Graenicher) 2 0.25% + + Lasioglossum nymphaearum (Cockerell) 3 0.37% + + Lasioglossum oblongum (Lovell) 3 0.37% + + Lasioglossum pectoral (Smith) 47 5.77% + + Lasioglossum pilosum (Smith) 15 1.84% + + Lasioglossum planatum (Lovell) 6 0.74% + + Lasioglossum quebecense (Crawford) 1 0.12% + Lasioglossum tegulare (Robertson) 1 0.12% + + Lasioglossum timothyi Gibbs 21 2.58% + Lasioglossum versans (Lovell) 1 0.12% + + Lasioglossum versatum (Robertson) 23 2.82% + + Lasioglossum vierecki Crawford 1 0.12% + Sphecodes banksii Lovell 1 0.12% + Sphecodes persimilis Lovell & Cockerell 1 0.12% + Sphecodes prosphorus Lovell & Cockerell 1 0.12% + + Sphecodes ranunculi Robertson 1 0.12% + Megachilidae *Anthidium oblongatum (Illiger) 2 0.25% + + Heriades carinata Cresson 2 0.25% + + Heriades leavitti Crawford 1 0.12% + Hoplitis producta (Cresson) 14 1.72% + + Hoplitis spoliata (Provancher) 1 0.12% + + Hoplitis truncate (Cresson) 17 2.09% + + Megachile gemula Cresson 4 0.49% + + Megachile latimanus Say 5 0.61% + + Megachile melanophaea Smith 14 1.72% + + Megachile mendica Cresson 2 0.25% + †Megachile mucida Cresson 1 0.12% Megachile relativa Cresson 2 0.25% + + Megachile texana Cresson 3 0.37% + Osmia atriventris Cresson 5 0.61% + + Osmia bucephala Cresson 3 0.37% + + Osmia collinsiae Robertson 1 0.12% + + Osmia distincta Cresson 1 0.12% + Osmia felti Cockerell 3 0.37% Osmia georgica Cresson 1 0.12% + Osmia pumila Cresson 5 0.61% + + Osmia virga Sandhouse 3 0.37% + Stelis labiate (Provancher) 4 0.49% + Northeastern Naturalist 386 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 abundance, although not significantly lower than the mowed-only or burned-only treatments. Bee richness was also significantly affected by management regime, with post-hoc analyses revealing that burned-then-mowed treatments contained significantly greater bee richness than all other treatments except the mowed-only Figure 2. Venn diagram depicting the number of unique wild bee species present within and shared among the Ossipee Pine Barrens (OPB; this study), the White Mountain National Forest (WMNF; Tucker and Rehan 2017a), and Strafford County, NH (Tucker and Rehan 2017b, 2018). The relative number of species collected at a site compared to all species collected at all sites is given as a percent in parentheses. Note that the Strafford County samples were accumulated over a longer period of time and throughout the active bee season, whereas, those from the OPB and WMNF are based on single-day collection events. Differences in unique species per surveyed area can in part be explained by unequal sampling effort. Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 387 treatment (χ2 = 14.38, df = 5, P = 0.002). Of the 59 bee species collected, 15% (9 species) were found in all management regimes and 55% (33 species) were exclusive to a single treatment. Discussion Bees of the OPB This study documents the first survey of wild bees in the OPB. We document the wild bee Megachile mucida for the first time in the state of New Hampshire. Although previously recorded in Maine, Massachusetts, Connecticut, and other nearby states, the species had yet to be documented in New Hampshire (Ascher and Pickering 2017, UNHC 2018). We also found 4 other species (Table 2) that while previously recorded for NH and neighboring states, have not been recorded in either Strafford County or the WMNF (Ascher and Pickering 2017, Tucker and Rehan 2017b, 2018). These findings demonstrate that there is still much need for research on the New Hampshire bee fauna (Koh et al. 2016). We also confirmed 3 introduced bee species in the OPB (see Table 2). Although not previously documented in the OPB, these introduced species have been recorded in the state and are commonly found throughout the Northeast (Droege 2015). The presence of Figure 3. Mean bee abundance and species richness relative to treatment regime (n = number of samples per treatment). Treatment significantly affected both bee abundance (dark grey bars) and richness (light grey bars) based on GLM results. Post-hoc analyses showed abundance, as well as richness, were significantly greatest in the burned and then mowed treatment (significant differences are indicated by letters in order from left to right). Northeastern Naturalist 388 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 introduced species in an already fragile ecosystem could indicate an impending decline in bee species dependent on this habitat (Gibbs and Sheffield 2009, Morales et al. 2013, Paini 2004). OPB bee similarity to nearby regions Despite being relatively close to the WMNF, we found OPB to have a very different wild bee composition, with 30 species not found in the WMNF (Tucker and Rehan 2017a). Further studies will be required to determine how species composition varies between these habitats due to floral resource or nesting site availability, elevation, or land management; other studies have found many unique and rare insect species in pine barrens (Bried et al. 2014, Grand and Mello 2004, Leuenberger et al. 2016, NatureServe 2017, Swengel 1998). Thorough and regionally specific studies are important to not only account for high levels of landscape specificity, but also ecological fluctuations and species turnover between years and seasons (Oertli et al. 2005, Rollin et al. 2015). The OPB bee community was similar to the Strafford County bee community: we found only 7 species in the OPB not recorded in Strafford County (Tucker and Rehan 2017b, 2018). Considering that the duration of the survey in Strafford County was much longer (3 years compared to 1 day), the difference in species composition is likely due to habitat differences. Some of these differences in species composition may also be attributed to the time of year sampled, as many species have limited flight periods, and OPB sampling would have missed many early spring, late summer, and fall species. We expect that additional surveying of the OPB will expand the species list, increasing both the number of unique species and species found in both habitats. Management regime effects In our survey, we found significantly greater bee abundance and richness in the burned-then-mowed sites than in the other sites. These results are consistent with bee surveys conducted in other fire-dependent habitats that found early successional habitats caused by burning supported larger bee communities (Campbell et al. 2018, Mola et al. 2018, Potts 2006, Rodriguez and Kouki 2015). Our findings that bee abundance in the mowed treatments did not significantly differ from those in our control sites were also similar to results from a comparable study on bees in the pine barrens of New York. In that study, Bried and Dillon (2012) found no significant differences in bee abundance and richness between fireprone sites that were mowed and treated with herbicides compared to those that were unmanaged and fire-prone. In addition to greater bee abundance and richness in burned-then-mowed sites, we found a high species–site exclusivity. While some of these species may truly only be associated with the specific management regime sampled, based on the high number of singleton species collected, it is likely some of the site exclusivity can be attributed to only collecting on a single day and the resulting small sample size. These results suggest that wild bees may benefit from periodic burning and infrequent mowing. Combined, these land-use management regimes may reduce the growth of non-flowering forbs and increase open ground for nesting sites. In addition to providing the first wild bee survey in Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 389 the OPB, this study highlights the need for long-term and repeated surveys of wild bees across the highly varied landscapes of New England and beyond. Acknowledgments We thank Joan Milam for assistance with bee collection and identification. We also thank all the participants of the NH Bee Bioblitz for their assistance with fieldwork and specimen processing. We thank the Nature Conservancy and NH Fish and Wildlife for permitting this event. Partial funding was provided by the New Hampshire Agricultural Experiment Station. This work was supported by the USDA National Institute of Food and Agriculture Hatch Project 1004515. Literature Cited Ascher, J.S., and J. Pickering. 2017. Discover Life bee species guide and world checklist (Hymenoptera: Apoidea: Anthophila). Available online at http://www.discoverlife.org/ mp/20q?guide=Apoidea_species. Bartomeus, I., J.S. Ascher, J. Gibbs, B.N. Danforth, D.L. Wagner, S.M. Hedtke, and R. Winfree. 2013. Historical changes in northeastern US bee pollinators related to shared ecological traits. Proceedings of the National Academy of Sciences 110:4656–4660. Bried, J.T., and A.M. Dillon. 2012. Bee diversity in Scrub Oak patches 2 years after mow and herbicide treatment. Insect Conservation and Diversity 5:23 7–243. Bried, J.T., W.A. Patterson, and N.A. Gifford. 2014. Why pine barrens restoration should favor barrens over pine. Restoration Ecology 22:442–446. Campbell, J.W., P.A. Vigueira, C.C. Viguiera, and C.H. Greenberg. 2018. The effects of repeated prescribed fire and thinning on bees, wasps, and other flower visitors in the understory and midstory of a temperate forest in North Carolina. Forest Science 64:299–306. Coelho, B.W.T. 2004. A review of the bee genus Augochlorella (Hymenoptera: Halictidae: Augochlorini). Systematic Entomology 29:282–323. Droege, S. 2015. The Very Handy Manual: How to Catch and Identify Bees and Manage a Collection. Native Bee Inventory and Monitoring Lab, USGS Patuxent Wildlife Research Center, Beltsville, MD. 65 pp. Gibbs, J. 2011. Revision of the metallic Lasioglossum (Dialictus) of eastern North America (Hymenoptera: Halictidae: Halictini). Zootaxa 3073:1–216. Gibbs, J., and C.S. Sheffield. 2009. Rapid range expansion of the Wool-Carder Bee, Anthidium manicatum (L.) (Hymenoptera: Megachilidae), in North America. Journal of the Kansas Entomological Society 82:21–29. Grand, J., and M. J. Mello. 2004. A multi-scale analysis of species–environment relationships: Rare moths in a Pitch Pine–Scrub Oak (Pinus rigida–Quercus ilicifolia) community. Biological Conservation 119:495–506. Grigarick, A.A., and L.A. Stange. 1968. The pollen-collecting bees of the Anthidiini of California (Hymenoptera: Megachilidae). Bulletin of the California Insect Survey 9:1–113. Grundel, R., R.P. Jean, K.J. Frohnapple, G.A. Glowacki, P.E. Scott, and N.B. Pavlovic. 2010. Floral and nesting resources, habitat structure, and fire influence bee distribution across an open–forest gradient. Ecological Applications 20:1678–1692. Horikoshi, M., and Y. Tang. 2015. ggfortify: Data visualization tools for statistical analysis results. v0.1.0. Available online at http://CRAN.R–project.org/package=ggfortify. Accessed 24 April 2018. Northeastern Naturalist 390 E.M. Tucker and S.M. Rehan 2019 Vol. 26, No. 2 Hothorn, T., F. Bretz, and P. Westfall. 2008. Simultaneous inference in general parametric models. Biometrical Journal 50:346–363. Howard, L.F., T.D. Lee, and R.T. Eckert. 2011. Forest community composition and dynamics of the Ossipee Pine Barrens, New Hampshire. The Journal of the Torrey Botanical Society 138:434–452. Koh, I., E.V. Lonsdorf, N.M. Williams, C. Brittain, R. Isaacs, J. Gibbs, and T.H. Ricketts. 2016. Modeling the status, trends, and impacts of wild bee abundance in the United States. Proceedings of the National Academy of Sciences 113:140–145. Laverty, T.M. and L.D. Harder. 1988. The bumble bees of eastern Canada. The Canadian Entomologist 120:965–987. Leuenberger, W., S. Bearer, J. Duchamp, S. Johnson, B. Leppo, P. McElhenny, and J. Larkin. 2016. A comparison of Lepidoptera communities inhabiting restored and latesuccessional Pitch Pine–Scrub Oak barrens in Pennsylvania. Natural Areas Journal 36:38–47. Michener, C.D. 2000. The Bees of the World. Johns Hopkins University Press, Baltimore, MD. 913 pp. Michener, C.D., R.J. McGinley, and B.N. Danforth. 1994. The Bee Genera of North and Central America (Hymenoptera: Apoidea). Smithsonian Institution Press, Washington, DC. 209 pp. Mitchell, T.B. 1960. Bees of the Eastern United States. North Carolina Agricultural Experiment Station, Raleigh, NC. 538 pp. Mitchell, T.B. 1962. Bees of the Eastern United States. North Carolina Agricultural Experiment Station, Raleigh, NC. 557 pp. Mitchell, T.B. 1980. A generic revision of the megachiline bees of the Western Hemisphere. Contribution from the Department of Entomology, North Carolina State University, Raleigh, NC. Mola, J.M., and N.M. Williams. 2018. Fire-induced change in floral abundance, density, and phenology benefits bumble bee foragers. Ecosphere 9:e02056. Morales, C.L., M.P. Arbetman, S.A. Cameron, and M.A. Aizen. 2013. Rapid ecological replacement of a native bumble bee by invasive species. Frontiers in Ecology and the Environment 11:529–534. NatureServe. 2017. NatureServe Explorer. Available online at http://explorer.natureserve. org/. Accessed 12 June 2018. Oertli, S., A. Mueller, and S. Dorn. 2005. Ecological and seasonal patterns in the diversity of a species-rich bee assemblage (Hymenoptera: Apoidea: Apiformes). European Journal of Entomology 102:53–63. Oliveros, J.C. 2007. VENNY. An interactive tool for comparing lists with Venn diagrams. Available online at http://bioinfogp.cnb.csic.es/tools/venny/index.html. Accessed 3 May 2018. Ordway, E. 1966. Systematics of the genus Augochlorella (Hymenoptera, Halictidae) north of Mexico. The University of Kansas Science Bulletin 46:509–624. Paini, D.R. 2004. Impact of the introduced Honey Bee (Apis mellifera) (Hymenoptera: Apidae) on native bees: A review. Austral Ecology 29:399–407. Patterson, T., R.S. Maxwell, G.L. Harley, J.S. Oliver, J.H. Speer, S. Collins, M. Downe, B. Gannon, L. Ma, and C. Raso. 2016. Climate–growth relationships of Pinus rigida (Mill.) at the species’ northern range limit, Acadia National Park, ME. Northeastern Naturalist 23:490–500. Plowright, R.C., and M.J. Pallett. 1978. A morphometric study of the taxonomic status of Bombus sandersoni (Hymenoptera: Apidae). The Canadian Entomologist 110:647–654. Northeastern Naturalist Vol. 26, No. 2 E.M. Tucker and S.M. Rehan 2019 391 Potts, S.G., T. Petanidou, S. Roberts, C. O’Toole, A. Hulbert, and P. Willmer. 2006. Plant– pollinator biodiversity and pollination services in a complex Mediterranean landscape. Biological Conservation 129:519–529. R Core Team. 2015. R: A language and environment for statistical computing. v3.1.3, R Foundation for Statistical Computing, Vienna, Austria. Available online at http:// www.R–project.org/. Accessed 24 April 2018. Rehan, S.M., and C.S. Sheffield. 2011. Morphological and molecular delineation of a new species in the Ceratina dupla species group (Hymenoptera: Apidae: Xylocopinae) of eastern North America. Zootaxa 2873:35–50. Rodríguez, A., and J. Kouki. 2015. Emulating natural disturbance in forest management enhances pollination services for dominant Vaccinium shrubs in boreal, pine-dominated forests. Forest Ecology and Management 350:1–12. Rollin, O., V. Bretagnolle, L. Fortel, L. Guilbaud, and M. Henry. 2015. Habitat, spatial, and temporal drivers of diversity patterns in a wild bee assemblage. Biodiversity and Conservation 24:1195–1214. Swengel, A.B. 1998. Effects of management on butterfly abundance in tallgrass prairie and pine barrens. Biological Conservation 83:77–89. The Nature Conservancy. 2018. Ossipee Pine Barrens: Freedom, Madison, Tamworth, Ossipee, NH. Available online at https://www.nature.org/ourinitiatives/regions/northamerica/ unitedstates/newhampshire/places–preserves/ossipee–pine–barrens.xml. Accessed 4 may 2018. Tucker, E.M., and S.M. Rehan. 2016. Wild bee pollination networks in northern New England. Journal of Insect Conservation 20:325–337. Tucker, E.M., and S.M. Rehan. 2017a. High-elevation refugia for Bombus terricola (Hymenoptera: Apidae): Conservation and wild bees of the White Mountain National Forest. Journal of Insect Science 17:1–10. Tucker, E.M., and S.M. Rehan. 2017b. Wild bee community assemblages across agricultural landscapes. Journal of Agricultural and Urban Entomology 33:1–28. Tucker, E.M., and S.M. Rehan. 2018. Farming for bees: Annual variation in pollinator populations across agricultural landscapes. Agriculture and Forest Entomology. DOI:10.1111/afe.12287. University of New Hampshire Insect Collection (UNHC). 2018. UNHC webPortal to insects and arthropods. Available online at https://unhcollection.unh.edu/database. Accessed 11 June 2018. Wheeler, A.G., Jr. 1991. Plant bugs of Quercus ilicifolia: myriads of mirids (Heteroptera) in Pitch Pine–Scrub Oak barrens. Journal of the New York Entomological Society 405–440. Wrobleski, D.W., and J.B. Kauffman. 2003. Initial effects of prescribed fire on morphology, abundance, and phenology of forbs in big sagebrush communities in southeastern Oregon. Restoration Ecology 11:82–90.