Northeastern Naturalist 502 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 22001188 NORTHEASTERN NATURALIST 2V5(o3l). :2550,2 N–5o1. 23 Bees (Hymenoptera: Apoidea) Foraging on American Cranberry in Massachusetts Anne L. Averill1,*, Martha M. Sylvia1, Noel Hahn1, and Andrea V. Couto1 Abstract - We studied diversity and abundance of bees foraging on Vaccinium macrocarpon (American Cranberry) flowers in southeastern Massachusetts during 8 bloom periods between 2007 and 2016. We documented 6 families, 20 genera, and 72 species of bees. Only Bombus (bumble bee) species were abundant in any of the collection years. Ways in which the upland areas surrounding cultivated cranberry beds and American Cranberry itself may be inhospitable for many small-bodied bee populations with limited flight ranges are reviewed. Bombus dominated (>90%) the collection. Comparison with observations in the first half of the 20th century, together with collections made during a less-intensive survey 25 years ago, suggest that 2 Bombus species are increasing in abundance, but that at least half of the Bombus species previously observed in this region have become rare or locally extinct. Introduction Mass bloom of cultivated Vaccinium macrocarpon Aiton (Ericaceae; American Cranberry) has been a major feature of Cape Cod and southeastern Massachusetts landscapes for many years. Beginning in the 1800s, growers planted seeds or cuttings from native vines (Eastwood 1856) in beds of coarse sand that overlay peat deposits that had formed in the poorly drained swales or kettle holes left by glaciers (Damman and French 1987). These plantings formed a dense, spreading mat, with solitary flowers appearing along 5–10-cm upright stems in June through July. In the Massachusetts counties of Plymouth, Bristol, and Barnstable, the most suitable wetland sites had been developed into cranberry farms by the early 1900s (Thomas 1990). Today, natural bogs supporting large stands of American Cranberry are rare (Cox and Walker 2012, Damman and French 1987). In such wild bogs, Bombus spp. (bumble bees) are the most numerous visitors to the succession of flowers blooming throughout the season (Heinrich 1979, Reader 1977). Bumble bees are also the most common native bees foraging on American Cranberry flowers in cultivated beds (Cane et al. 1996, Evans and Spivak 2006, MacKenzie and Averill 1995). The observations of Henry J. Franklin (1913), who completed a comprehensive revision of North American bumble bees and studied all facets of cranberry cultivation for nearly half a century (1909–1953), provides a fortunate historical context for our work. Franklin (1950:75) observed that bumble bees were sometimes in “incredible” abundance on cranberry farms. On the other hand, native non-Bombus 1University of Massachusetts – Amherst, Department of Environmental Conservation, 160 Holdsworth Way, Amherst, MA 01003. *Corresponding author - averill@eco.umass.edu. Manuscript Editor: Kent McFarland Northeastern Naturalist Vol. 25, No. 3 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 503 bees received little attention, and were mentioned as “more or less common” on cranberry bogs (Franklin 1950:76). In 1990–1991, our limited study found a low abundance (8.3% of individuals in the total collection) of non-Bombus bees (Mackenzie and Averill 1995). In the present note, our goal is to update the inventory of all bee fauna foraging on American Cranberry in Massachusetts and to briefly compare Bombus species diversity to that reported by H.J. Franklin and, more recently, by MacKenzie and Averill (1995). Methods To create a baseline of bee fauna visiting American Cranberry, we processed and identified bees that were collected during studies on bee health, e.g., examining parasite loads and anthropogenic challenges (such as pesticide exposure or land-use change). Collections were carried out over 8 years (2007–2009, 2011–2013, 2014, 2016) and encompassed ~500 visits to 49 different cultivated bogs under conventional management (Fig. 1). Organically managed bogs were too few to comprise a representative group. Sites varied in size (mean = 7.5 ha, min–max = 0.5–61 ha) and varied in landscape contexts, including those characterized as forest, urban, and agricultural. At 4 bogs, introduced colonies of the commercial bumble bee B. impatiens were regularly used, and at 7 others, they were occasionally used. We discuss the implications of this practice below. We usually visited each study site 3 times during bloom from mid-June through mid-July. Upon each bog visit, we either (1) collected wild bees from an American Cranberry flower by net or in vials during a 10–20-minute transect starting on the edge and walking slowly across the bog (2007–2009, 2013, 2014, 2016), or (2) collected 15 Bombus and 15 non-Bombus bees (2011, 2012). Apis mellifera L. (Honey Bee) is not included since abundance data were not collected in all years. Further, the huge majority of Honey Bee individuals originate from managed hives that are installed at bog sites at bloom and then removed. Results and Discussion We collected a total of 8128 bees. Each species and its corresponding number of collected individuals summed across years are listed in Table 1. This collection represents 6 families, 20 genera, and 72 species and comprises about 20% of the 377 known bees in Massachusetts (Goldstein and Asher 2016). Numerically, the collection was strikingly dominated by 5 species of bumble bee (>90% of total bees). Bombus impatiens made up more than half (56.7%) of the total number of bees, while Bombus bimaculatus (18.2%), Bombus perplexus (7.8%), Bombus griseocollis (7.3%), and Bombus vagans (3.8%) were less common. Several species were rare: 5 Bombus fervidus, 3 Bombus terricola, 4 Bombus sandersoni, 2 Bombus citrinus (a social parasite) and 1 Bombus affinis (collected in 2008) were recorded. Our counts reflect worker and male numbers of eusocial species during our limited sampling period of American Cranberry bloom. Even if a spring begins with the same number of founding queens, abundance in the collection may be greatly Northeastern Naturalist 504 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 Vol. 25, No. 3 affected by bee species phenology and colony size; for example, queens may become active much earlier in time and form large colonies that peak during cranberry Figure 1. Map showing location of sites for bee collections. Not all sites were sampled in all years of the study. Northeastern Naturalist Vol. 25, No. 3 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 505 Table 1. Totals for each bee species that was hand-collected from cultivated American Cranberry flowers in southeastern Massachusetts and Cape Cod, compiled from 8 of the years between 2007 and 2016. [Table continued on following page.] Family Subfamily Species n Andrenidae Andreninae Andrena bradleyi Viereck 2 Andrena carlini Cockerell 18 Andrena ceanothi Viereck 1 Andrena crataegi Robertson 3 Andrena cressonii Robertson 2 Andrena imitatrix Cresson 2 Andrena nasonii Robertson 1 Andrena nivalis Smith 25 Andrena nuda Robertson 5 Andrena perplexa Smith, 4 Andrena rugosa Robertson 1 Andrena spiraeana Robertson 1 Andrena vicina Smith 56 Andrena wilkella (Kirby) 7 Apidae Apinae Bombus affinis Cresson 1 Bombus bimaculatus Cresson 1476 Bombus citrinus (Smith) 2 Bombus fervidus (Fabricius) 5 Bombus griseocollis (DeGeer) 596 Bombus impatiens Cresson 4607 Bombus perplexus Cresson 637 Bombus sandersoni Franklin 3 Bombus terricola Kirby 3 Bombus vagans vagans Smith 316 Habropoda laboriosa (Fabricius) 1 Anthophorine Melissodes desponsa Smith 1 Nomadinae Nomada maculata Cresson 2 Nomada rodecki Mitchell 3 Xylocopinae Xylocopa virginica (L.) 23 Ceratina calcarata Robertson 2 Colletidae Colletinae Colletes thoracicus Smith 1 Colletes validus Cresson 1 Hylaeinae Hylaeus modestus Say 1 Halictidae Halictinae Agapostemon sericeus (Forster) 11 Agapostemon texanus Cresson 13 Agapostemon virescens (Fabricius) 6 Augochlora pura (Say) 2 Augochlorella aurata (Smith) 16 Augochloropsis metallica (Fabricius) 44 Halictus confusus Smith 11 Halictus ligatus Say 5 Halictus rubicundus (Christ) 18 Lasioglossum acuminatum McGinley 20 Lasioglossum bruneri (Crawford) 1 Lasioglossum cinctipes (Provancher) 10 Lasioglossum coriaceum (Smith) 6 Lasioglossum cressonii (Robertson) 2 Lasioglossum ellisiae (Sandhouse) 2 Northeastern Naturalist 506 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 Vol. 25, No. 3 bloom (e.g., B. impatiens), in comparison to queens that emerge later and/or may form smaller colonies (e.g., B. vagans) (Plath 1934, Williams et al. 2014). Non-Bombus wild bees made up 5.6% of the total number of bees counted. Of the 62 species, more than half was represented by 1 or 2 individuals, and no species could be considered abundant. As percentages of each family to the entire collection, this non-Bombus segment of the collection was comprised of Andrenidae (1.6%), non-Bombus Apidae (0.4%), Colletidae (0.04%), Halictidae (2.6%), Megachilidae (0.6%), and Melittidae (0.8%). The family Andrenidae was represented by 14 species of Andrena. Individuals of Andrena carlini, Andrena nivalis, and particularly, Andrena vicina were most often recorded for that family. Of the non-Bombus Apidae, we recorded 6 species in 5 genera. We found 3 species in Colletidae, each represented by a single specimen. All 27 species in the family Halictidae were in the large subfamily Halictinae. The most species-rich genus was Lasioglossum, half of which were metallic Lasioglossum (Dialictus). For both Halictus and Agapostemon, 3 of the 4 northeastern species in each genus were collected, as were all 3 of the augochlorine species (Augochlora pura, Augochlorella aurata, and Augochloropsis metallica). Augochloropsis metallica was collected in all but 1 of the sampling years. We recorded 11 species of Megachilidae, most of which were Megachile or Osmia species. A single Melittidae species, the Vaccinium specialist Melitta americana, was the most abundant (67 individuals) of all of the non-Bombus species. Table 1, continued. Family Subfamily Species n Lasioglossum georgeickworti Gibbs 2 Lasioglossum leucozonium (Schrank) 2 Lasioglossum lineatulum (Crawford) 3 Lasioglossum oblongum (Lovell) 1 Lasioglossum obscurum (Robertson) 1 Lasioglossum paraforbesii McGinley 1 Lasioglossum pectorale (Smith) 7 Lasioglossum pilosum (Smith) 2 Lasioglossum rohweri (Ellis) 2 Lasioglossum smilacinae (Robertson) 1 Lasioglossum trigeminum Gibbs 1 Lasioglossum versatum (Robertson) 17 Megachilidae Megachilinae Hoplitis truncata (Cresson) 1 Megachile addenda Cresson 9 Megachile gemula Cresson 4 Megachile mendica Cresson 9 Megachile texana Cresson 1 Osmia atriventris Cresson 1 Osmia cornifrons (Radoszkowski) 1 Osmia inspergens Lovell and Cockerell 1 Osmia pumila Cresson 1 Osmia virga Sandhouse 17 Coelioxys rufitarsis Smith 1 Northeastern Naturalist Vol. 25, No. 3 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 507 In addition to M. americana, according to Goldstein and Asher (2016), additional Ericaceae specialists that are regionally associated with Vaccinium include Andrena bradleyi, Osmia virga, Megachile addenda, and Colletes validus, all of which were present in our collections. On the other hand, another Vaccinium specialist, Andrena carolina Viereck, was never caught, probably because it is such an early season species, appearing well before American Cranberry bloom (Goldstein and Asher 2016). While we may anticipate that 25% of this Massachusetts collection would exhibit parasitic behavior (Goldstein and Asher 2016), parasitic species were rare in our collections (~7% of all species), and 8 individuals in 4 species were recorded: B. citrinus, known to parasitize B. impatiens, B.vagans, and B. bimaculatus (Williams et al. 2014); Nomada maculata, which was associated at 2 sites with Andrena carlini (possibly a host); Nomada rodecki, which we found associated at 3 sites with M. americana (this association is also reported in Goldstein and Asher 2016); and a single Coelioxys rufitarsis, which we found associated with Megachile mendica (possibly a host). Three species known or suspected to be exotic were collected: Lasioglossum leucozonium, Andrena wilkella, and Osmia cornifrons. Several new county records were found for Bristol County (Andrena perplexa, A. wilkella, and Ceratina calcarata) and for Plymouth County (Lasioglossum ellisiae, Lasioglossum lineatulum, Lasioglossum paraforbesii, Lasioglossum rohweri, and Lasioglossum trigeminum). While our survey is consistent with historical records showing that bumble bees are dominant members of the bee community in cranberry bogs, it appears that at least half of the Bombus species previously observed in the region have become rare or locally extinct. In North America, some species, including B. fervidus and Bombus pensylvanicus (DeGeer), have undergone range reduction and decline in abundance gradually over decades, while others, including B. terricola and Bombus affinis have undergone population collapse in the past 15 years (Cameron et al. 2011). The US Fish and Wildlife Service finalized the listing of B. affinis as an endangered species in 2017 (USFWS 2017). We also observed these long-term or precipitous patterns of loss in our study region. In 1950, B. pensylvanicus and Bombus ternarius Say were regularly found in southeastern Massachusetts (Franklin 1950), but none were collected in the 1990s survey (Mackenzie and Averill 1995) or in the present survey. Bombus fervidus (status = vulnerable; IUCN Red List of Threatened Species 2017), which Franklin (1950) called one of the most common species in the cranberry-growing region, was not collected in the 1990s, and we recorded only 5 individuals in the present survey. Perhaps the transition away from farms, pasture maintenance, and haying operations, currently 8% of Massachusetts land compared to 30–40% in Franklin’s time (USDA Census of Agriculture Historical Archive), has been an impactful landuse change. Franklin (1913:393, 405) noted that B. fervidus nests “are the ones most commonly found by the New England farmer during haying season” and for B. pensylvanicus, he wrote that all of the nests which “I have known about were in open grassland”. Franklin (1913) called Bombus sandersoni (Franklin) one of Northeastern Naturalist 508 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 Vol. 25, No. 3 the most common bumble bee species within its range, which includes Massachusetts, but did not mention it in his cranberry-related writings, perhaps because it is associated with forest sites (Williams et al. 2014). Recent assessment of its persistence has been problematic (Goldstein and Asher 2016, IUCN 2017). No B. sandersoni were collected in an earlier survey (Mackenzie and Averill 1995), but a single capture was recorded in each of 3 years in the present study. On the other hand, while Bombus affinis and B. terricola were well represented in our 1990s survey, we found only 1 and 3 specimens, respectively, similar to findings of recent population collapse reported throughout North America (Cameron et al. 2011). The bees were collected only in 2008–2009 on 2 isolated farms, one in northern Cape Cod and another within Myles Standish State Forest. Compared to proportions of species seen in the 1990s (Mackenzie and Averill 1995), B. impatiens and B. griseocollis appear to be increasing, and this trend is observed elsewhere in the country (Bartomeus et al. 2013, IUCN 2017). These 2 species are known to thrive in urban areas (Williams et al. 2014), and our study region lies within the heavily urbanized Boston–Washington corridor. A confounding factor in the present study is that at 22% of the sites, growers (for varying numbers of years) introduced colonies of commercially cultured B. impatiens for pollination. The cultured individuals cannot be distinguished from wild B. impatiens, so some captures may have originated from active commercial hives. However, our recent genetic analyses of B. impatiens foraging in our study area (Suni et al. 2017) provide no support for the hypothesis that these commercial hives are measurably contributing to an upward population trend. First, even with active commercial B. impatiens hives present, we rarely detected foragers on the bogs, perhaps because the probability of capture is swamped by very large wild bee populations, or perhaps because these commercial bees were foraging elsewhere. Second, there was no widespread introgression of alleles from commercial bumble bees to wild bees and that the commercial bees have not established in the bogs (Suni et al. 2017). Expanded studies are required to document the impact of importing commercial bees under varying conditions. Overall, many consecutive years of survey are required to be truly certain if a species is declining (Goldstein and Asher 2016) or increasing, and why these changes are occurring. Change in population numbers, even for common bee species, is one of the biggest knowledge gaps in consideration of bee population status (Goulson et al. 2015). Regarding the low abundance of non-Bombus bees on cultivated bogs, although the region’s sandy soils may provide appropriate nest sites for many species, we suspect that the upland areas surrounding cultivated cranberry beds and American Cranberry plants themselves may be inhospitable for many small-bodied bees that are likely to have more limited flight ranges than larger bees (Greenleaf et al. 2007). Regarding landscape conditions, cranberry beds are within the coastal sand plain region of MA, which is striking in its lack of diverse floral resources. These off-bog upland areas typically consist of Pinus rigida Mill. (Pitch Pine)/Quercus ilicifolia Wangenh (Bear Oak) forests and have been called “waste land” (Franklin 1950:75). Sparse spring pollen sources may Northeastern Naturalist Vol. 25, No. 3 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. Couto 2018 509 challenge bees with early-onset flight seasons, particularly some Andrenidae and Halictidae. On-bog conditions may be equally challenging. A critical concern is pesticide impact. For native bees, exposure and sensitivity to pesticides may be different from those determined for the Honey Bee, the test species most often used in pesticide risk-assessment bioassays (Arena and Sgolastra 2014). Such bioassays are the basis for labeled use patterns of compounds applied during bloom (Fischer and Moriarty 2014). Second, unlike other crops, for example, ecologically managed tree fruit or nut orchards (e.g., Saunders et al. 2013), American Cranberry bogs have no understory of clovers, weedy Asteraceae, or other alternative pollen sources. Third, although the thick sandy deposits on cultivated bogs could support ground-nesting species, nests likely would be eliminated by floods applied for harvest and winter protection. Finally, American Cranberry flowers do not produce easy-access pollen since it is both hidden and firmly held within the anthers. Only some bees are effective collectors, and do so by shaking the pollen loose from the anthers (Buchman 1983, De Luca and Vallejo-Marin 2013). Such “buzz pollination” is well described in the bumble bees, and is demonstrated by a number of other species, e.g., Agapostemon, Augochloropsis, and the small family Melittidae (Buchman 1983). We observe that our methods and resulting inventory differ considerably from 2 other bee collections in American Cranberry bogs. We limited our sampling to bees that were hand-collected directly from American Cranberry flowers during the month of bloom. In contrast, in a Wisconsin study, Gaines Day (2013) deployed bowl traps (Cane et al. 2000) in bogs from May to August and reported large collections of non-Bombus individuals (particularly Agapostemon and Lasioglossum) and species richness over 2.4 times (182 bee species) that of our Massachusetts study. Additionally, 7% of total bees captured in the Wisconsin Gaines Day (2013) study were Bombus species, in comparison to ~90% of our captures. A good deal of this latter discrepancy can be explained by the fact that pan-trapping is not a good method for estimating the abundance of foraging bumble bees (Baum and Wallen 2011, Bushman and Drummond 2015). In a second cranberry study, Loose et al. (2005) employed flight-intercept, sticky, and malaise traps deployed at various heights over the entire season across southeastern Massachusetts bogs and into adjoining habitats. Fifty species in 17 genera were reported, and the collection was dominated by Megachile addenda (>80%), while Bombus spp. made up 13.6% of captures. We again suspect that abundance of this latter group is underestimated by the trapping methodology. Finally, documentation of a species in the bog system (or, as in our study, that it is captured on the flower) does not confirm its utilization of available floral resources or its role as a crop pollinator. Indeed, while bumble bees are well-known pollen collectors and pollinators of American Cranberry (Broussard et al. 2011, Cane and Schiffhauer 2003, Franklin 1950, MacKenzie 1994, Ratti et al. 2008), these traits are poorly known for most of the non-Bombus species that either we or the above researchers collected and thus, these questions will require further study. Northeastern Naturalist 510 A.L. Averill, M.M. Sylvia, N. Hahn, and A.V. 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