Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 47 2014 NORTHEASTERN NATURALIST 21(1):47–71 Bees (Hymenoptera: Apoidea: Anthophila) of Gardiners Island, Suffolk County, New York John S. Ascher1,2,*, Sarah Kornbluth1,3, and Robert G. Goelet1 Abstract - We present a survey of the bee fauna of Gardiners Island, Suffolk County, NY. The study focuses on more than 10,000 bee specimens collected by net during 1976, 1977, and 2005–2011. These surveys recorded 154 bee species and morphospecies in 30 genera. Bee species associated with fields, beaches, and woody plants were well represented in the sample, whereas those requiring herbaceous understory flowers were relatively few. Notable finds include several species with southern affinities that are otherwise scarce in New York State and paratypes of the social parasite Lasioglossum (Dialictus) rozeni. The newly discovered male of Sphecodes johnsonii is diagnosed and imaged. Two Nomada morphospecies may be undescribed. A single worker of the northern species Bombus (Cullumanobombus) rufocinctus provides the first coastal record for New York State. New records of Dianthidium (D.) simile from nearby Long Island and of Pseudoanthidium (P.) nanum from New York City (and nearby New Jersey) increase the number of bee genera and described species recorded from New York State to 45 and 447, respectively. Phenological data may provide new evidence for host-parasite associations. The bee diversity of Gardiners Island is discussed in relation to the bee fauna of the northeastern US, especially records available for other Atlantic coastal islands in the region. Introduction In recent years, there has been growing interest in documenting the status of North American bees due in part to well-publicized threats to these pollinators (NAS 2007). Most species known historically from eastern North America persisted to 1990–2009 despite severe and well-documented declines in certain taxa (Bartomeus et al. 2013, Colla et al. 2012), but comprehensive publications on local bee faunas in the region remain few. There are not many published studies on the occurrence of bee species in New York State (e.g., Leonard 1928; state records in Mitchell 1960; see also Giles and Ascher 2006), but J.S. Ascher (unpubl. data) has compiled a list of 447 described species (mapped online at Discover Life [Ascher and Pickering 2013]) of which at least 21 are exotic. Although online state lists of bee species known from New York and the New England states are now relatively complete, precise distributional patterns remain poorly documented. The bee fauna of eastern Suffolk County, Long Island (LI) was surveyed by the famed naturalist Roy Latham in the first half of the 20th century, including large 1Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 81st Street, New York, NY 10024. 2Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543. 3Department of Biological Sciences, Rutgers University, 101 Warren Street, Newark, NJ 07102. *Corresponding author - dbsajs@nus.edu.sg. Manuscript Editor: Kent McFarland Northeastern Naturalist 48 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 collections from his home at Orient, the eastern-most town on Long Island’s North Fork, located about 10 mi WNW of Gardiners Island (GI). Some of Latham’s bee specimen records have been documented in published revisions, but most remain unpublished, although they are now being digitized through specimen databasing at Cornell University (Schuh et al. 2010). The Latham collection includes records of several coastal bee species at or near their northeastern distributional limits. William T. Davis and others also made important historical collections from LI at Wading River in Suffolk County on the North Shore during 1914–1922. Specimens of Dianthidium (D.) simile (Cresson) from that locality in the University of California, Riverside Collection extend the range of this genus and species to New York. Bee species totals for all 43 genera then known to occur in New York State were summarized by Giles and Ascher (2006). Subsequently, adventive populations of the Palearctic genus Pseudoanthidium, were discovered in the New World by S. Kornbluth in northern New Jersey and identified as Pseudoanthidium nanum (Mocsáry) with reference to the work of M. Schwarz (Aguib et al. 2010). The species has also been recorded from New York City (K.C. Matteson, Miami University, Oxford, OH, pers. comm) and Maryland (Droege and Shapiro 2011). Thus, 45 bee genera have now been recorded from New York State, of which 43 include at least one native species, whereas Apis and Pseudoanthidium are each represented by one exotic species. Here we summarize extensive net collections of bees from GI during 2005–2011 and a smaller number of earlier collections made there, mostly during 1976–1977. We interpret the resulting fauna in comparison to other historical and recent collections available from the northeastern US. Collections from GI are of particular interest, as this privately owned island has been spared the intensive development typical of western LI within and adjacent to New York City (NYC). The present study complements published faunal surveys for urban NYC (Matteson et al. 2008), nearby suburbs (Fetridge et al. 2008), and natural areas (Giles and Ascher 2006). Data from GI are also intended for comparison with bee surveys now in progress for multiple Atlantic coastal islands in the region including Martha’s Vineyard (P.Z. Goldstein, National Museum of Natural History, Washngton, DC, unpubl. data), Nantucket (P.Z. Goldstein, unpubl. data), the Elizabeth Islands (G.I. Stage, Stafford Springs, CT, unpubl. data) of Massachusetts, Block Island of Rhode Island, and a statewide bee survey of Connecticut (by J.S. Ascher et al., unpubl. data). Research Objectives This survey had four primary goals: 1) to assemble a species list of Gardiners Island bees for future comparison with bee surveys of other Atlantic coastal islands; 2) to contribute to a growing database of specimen records for regional bees used to investigate phenology (Bartomeus et al. 2011), status (Bartomeus et al. 2013), and other aspects of bee distribution and ecology; 3) to investigate host-parasite relationships of bees and other aspects of their life history; and 4) to enhance representation of bees in the collection and database of the American Museum of Natural History (AMNH). Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 49 Study Site Gardiners Island (1343 ha [3318 acres]) is located between the two peninsulas of eastern LI, northwest of Montauk and under the jurisdiction of the town of East Hampton (41°03'–41°07'N, 72°05'–72°08'W; Fig. 1). It forms the boundary between Gardiners Bay and Block Island Sound, east of Shelter Island and the entrance to Peconic Bay, and is six miles long and three miles wide. Nearby islands include Plum Island 4.5 mi NNW, Little Gull and Great Gull Islands 5.3 mi N, Fishers Island 10 mi NNE (these are all in Suffolk County, NY), and Block Island 25 mi ENE in Washington County, RI. These islands are all part of the Outer Lands, a terminal moraine archipelagic region off the southern coast of New England. This region of Massachusetts, Rhode Island, and New York comprises the peninsula of Cape Cod and the islands of Martha's Vineyard, the Elizabeth Islands, Nantucket, Block Island, and Long Island, as well as surrounding islets. The original biota of GI is unknown, but we do know that American Indians occupied it. Large changes in habitat resulted from a long history of human occupation and agricultural use of the island after 1639, when it was deeded to Lion Gardiner by King Charles I. In recent times, it has had few inhabitants and limited visitation and has been spared the residential and commercial development prevalent in the region. The island has relict Bostwick Forest with notably old and large Quercus alba L. (White Oak), other woodlands, and several hundred acres of fields maintained by seasonal mowing. Habitats significant to bees include sandy, gravel, or cobble beaches extending around the island and associated salt marshes and brackish ponds lined with Hibiscus moscheutos Welw. ex Hiem (Rose Mallow). Among the beach flora, Solidago sempervirens L. (Seaside Goldenrod) is a particularly important bee host plant. Cliffs composed of glacial till in the northern part of the island rise as high as about 120 feet and are the habitat of flowers such as Euthamia. Figure 1. Map of Gardiners Island area including Fishers, Block, and Great Gull islands. Source: http://www.demis.nl. Northeastern Naturalist 50 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 “The flora of Gardiners Island” by Hehre (1977; updating Burnham and Latham 1914) provides a site description and lists 403 plant taxa. The author noted two factors that have likely reduced floral resources available to bees: 1) a large deer population resulting in “high browse-lines at the edge of woodlands”, and 2) large populations of Smilax rotundifolia L. (Common Greenbrier) as well as Vitis labrusca L. (Fox Grape), a native but aggressive species that “eliminated most other vegetation from a large proportion of the Bostwick Forest.” Important resources for bees from spring to early summer include Vaccinium (blueberries; V. macrocarpon Aiton [Cranberry] is not recorded), four species of Rhus (sumac), and various Rosaceae. In midsummer, Cirsium (thistles) and introduced Trifolium (clovers) in fields provide food for bees, whereas four recorded species of Solidago sensu lato (goldenrods, including species now placed in genus Euthamia) and other composites are the most important host plants blooming in late summer and autumn. The only pines on the island are planted stands, whereas pine barren habitats provide important bee habitat on Long Island and on Martha’s Vineyard. Willows are poorly represented, with only Salix nigra Marsh. (Black Willow) recorded. Methods Sampling methods and schedule Bees were collected by net, most while visiting flowers, by R.G. Goelet on GI on 8 days during 1976 (21 Jun–18 Jul), 8 days in 1977 (22 Apr–30 May and 15 Jul), 15 and 21 Jun 1986, 1 Oct 2001, 56 days in 2005 (1 May–7 Oct), 59 days in 2006 (13 Apr–9 Oct), 63 days in 2007 (28 April–21 Oct), 32 days in 2008 (24 May–27 Oct), 27 days in 2009 (25 Apr–14 Oct), 20 days in 2010 (1 May–2 Sep), and 15 days in 2011 (1 May–7 Aug). In addition, J.S. Ascher and J.G. Rozen, Jr. collected together with R.G. Goelet on 4–5 Aug 2007. In all, bees were collected on 292 days (290 of them by R.G. Goelet alone), and were searched for on additional days when weather proved too inclement to make a collection. Most intensive fieldwork was conducted on days with predominantly sunny skies and warm temperatures. Collecting activity on the island was opportunistic in response to the presence of flowering plants and bee visitors. The primary collector (R.G. Goelet) is not a bee specialist. He made an effort to obtain a maximum diversity of aculeate Hymenoptera, including wasps (relevant because without searching for these, small cleptoparasitic bees and others might be overlooked), on a variety of plants. Specimen processing and databasing Specimens were identified to species by J.S. Ascher with assistance from colleagues. The more difficult metallic Lasioglossum (Dialictus) were determined by J. Gibbs, and many female Ceratina calcarata and dupla were separated by S. Rehan. Vouchers are deposited in the collection of the American Museum of Natural History (AMNH). Duplicate specimens were sent to the Canadian Centre for DNA Barcoding, Guelph, Canada for DNA sequencing of cytochrome c oxidase subunit 1 as a contribution to the Bee Barcode of Life Project. Most bees collected were fully processed and entered in the AMNH Division of Invertebrate Zoology Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 51 Database (Schuh et al. 2010), but some samples of very common species such as Halictus ligatus, Bombus impatiens, and B. griseocollis from 2008–2010 were not mounted, and some duplicate specimens were dispersed for DNA sequencing or other purposes prior to being databased. Thus, the reported totals are slightly lower than the actual totals of specimens collected for many species, but much lower only for a few of the most common species, i.e., those already present in long series so deliberately not pinned due to insufficient resources. Comparative data on the North American bee fauna as a whole and on the faunas of New York State (NY), particular areas within NY, and other states in the northeastern US were compiled by J.S. Ascher based on the literature and historical insect collections, especially those housed at the AMNH; Cornell University (CUIC); New York State Museum (NYSM); National Museum of Natural History (NMNH); University of Connecticut Insect Collections, Storrs (UCMS); Rutgers University Arthropod Collection (RUAC); University of California Riverside (UCRC); and Parker Gambino’s personal collection (affiliated with the AMNH). Recent collections from southern NY and nearby Connecticut were available for comparison, including voucher specimens from recent ecological studies (e.g., Fetridge et al. 2008, Giles and Ascher 2006, Matteson et al. 2008). All discussion of the past and present status and life histories of bee species found at GI are based, in part, on these historical and recent collections and the literature, in addition to the collections obtained during the survey of GI. Specimen records in the AMNH Division of Invertebrate Zoology database (Schuh et al. 2010) and others captured using Arthropod Easy Capture (2013) software provided context for GI records and included 47,734 bees from New York State, of which 1903 were from greater Long Island exclusive of GI (but including Kings and Queens counties of New York City); 19,552 from Connecticut; and 7432 from New Jersey. Results and Discussion During the survey period, 10,674 bee specimens from GI, representing 154 species and morphospecies of which 150 are native and five are exotic (Appendix 1), were collected, mounted, identified, and databased. These belong to five families (no melittids were found) and 30 of the 45 bee genera recorded from New York. The phenological patterns of the bee community of GI (Fig. 2) is similar to that seen in other studies from this region (Giles and Ascher 2006), with large numbers of Andrena active in the early spring along with queens of eusocial species, while males and workers of eusocial species were more numerous in late summer and fall. Differences include a wider variety and greater abundance of late-summer and fall Andrena and Colletes and their respective Nomada and Epeolus cleptoparasites (Fig. 3). These species are notably diverse and numerous at sandy coastal sites, including islands, with extensive seasonal bloom of composites. Figure 2 (following 2 pages). Phenology of all identified species except Apis mellifera and Hylaeus punctatus, with the date range expressed for males above in black and females below in grey. Numbers in parentheses are totals of male and female specimens databased. Northeastern Naturalist 52 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 53 Northeastern Naturalist 54 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Summary of findings by taxa Colletidae. We collected eight species of Colletes, a high number for the northeastern United States, reflecting availability of appropriate host plants including composites in conjunction with sandy nesting substrates. Colletes speculiferus (= mitchelli Stephen; see Kuhlmann and Ascher 2011) is a late-flying specialist of Solidago sensu lato, particularly S. sempervirens growing on coastal sand dunes. The very similar C. americanus Cresson and another member of the americanus species group, the regionally scarce C. solidaginus, were also collected on Solidago as were, in larger numbers, the widely distributed C. compactus compactus Cresson and C. simulans armatus Patton. Colletes nudus has been found in New York only on Long Island (including Brooklyn, K.C. Matteson, unpubl. data), and the 54 recorded from GI are the largest known collection from the northeastern US. Males were numerous on 4–5 Aug 2007 visiting flowers of Rhus (sumac), likely a preferred host plant, and also Melilotus alba Medik. (White Sweetclover), likely an incidental nectar source. Our sample of Hylaeus includes four native species of which Hylaeus (Prosopis) cf. nelumbonis (Robertson), a relatively large species with infumate wings, is most noteworthy. The two male specimens, one from 17 Jul 1976 and the other from 5 Aug 2007, have a black first tergum and therefore appear to be Hylaeus schwarzii (Cockerell), but lack the conspicuous basal elevations on the third and fourth sterna of that species and instead have a rugose pleuron and metanotum, characters diagnostic of H. nelumbonis (Mitchell 1960). Hylaeus nelumbonis appears to be associated with wetlands with the few NY records coming from sites such as Montezuma National Wildlife Refuge at the north end of Cayuga Lake in the Finger Lakes Region (J.S. Ascher, unpubl. data). The host plants of this bee on GI are unknown, as neither Nelumbo nor Nymphaea from which H. nelumbonis has been collected (Mitchell 1960) is listed in the GI flora (Hehre 1977). Absence of H. schwarzii from our GI collections is noteworthy, as this species but not H. nelumbonis is recorded from the Elizabeth Islands (G.I. Stage et al., unpubl. data) and Martha’s Vineyard (P.Z. Goldstein and J.S. Ascher, unpubl. data). The introduced species Hylaeus (Spatulariella) punctatus was only recently detected in eastern North America but is now widely distributed (Sheffield et al. 2011). It was known previously in NY only from the New York City area (Ascher et al. 2006). Figure 3. Phenology of suggested host-parasite relationships with log-scaled abundance. Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 55 Halictidae. All three augochlorine species known from NY are numerous on GI, with ground-nesting Augochlorella aurata the most abundant, whereas the woodnesting Augochlora pura is the most abundant augochlorine species in New York City (Matteson et al. 2008). The two most common species of Agapostemon in NY, A. virescens and A. sericeus, were very numerous, and 8 A. splendens and 3 A. texanus were also collected. Of these, A. splendens is associated with sandy nesting substrates (Roberts 1969). The 10 species of the cleptoparasitic genus Sphecodes collected included S. ranunculi and also S. aroniae, a more southerly distributed species of the ranunculi species group for which Proteraner Robertson is an available genus-group name. Among species of the cressonii group of Sphecodes, for which Sphecodium Robertson is an available name, both sexes of S. johnsonii were found, the male of which is newly recognized. Males of S. johnsonii (Fig. 4) resemble males of S. davisii in having the scutum rugose along the anterior margin with the rest shining and punctate, densely punctate metasomal sterna 3-5, and relatively large body size, but the former has an all black metasoma whereas the latter has metasomal terga 1-3 red. Structural characters differentiating S. johnsonii from S. davisii in both sexes include dorso-lateral angle of pronotum forming a right angle in dorsal view as opposed to obtusely angled and the first flagellomere considerably broader than long (about 1/2 in the male and 2/3 in the female) as opposed to about as broad as long. Both sexes of S. johnsonii are very similar to S. galerus Lovell and Cockerell, but the latter is smaller (length of females 6 mm for galerus and 9–10 mm for johnsonii according to Mitchell [1960]). Sphecodes johnsonii males superficially resemble those of certain species in the dichrous group (separated at couplet 6 in the key to Sphecodes males in Mitchell [1960]) but differ from them in lacking pilose, semicircular facets on the antennal flagellum below. All four Halictus species found in the eastern United States were collected, of which H. ligatus was extremely numerous (895 specimens databased). Subgenus Lasioglossum was represented by L. coriaceum, widely distributed across the northeastern US, and by L. fuscipenne, found in NY mostly in the eastern and coastal portions of the state (McGinley 1986). Absence of L. acuminatum was surprising as this species is recorded from nearby Orient (McGinley 1986) and is common on Martha’s Vineyard (P.Z. Goldstein and J.S. Ascher, unpub. data). The widely distributed, pollen-generalist species Lasioglossum quebecense was collected. It has traditionally been classified as an Evylaeus (Mitchell 1960) but is now included in an expanded subgenus Sphecodogastra (Gibbs et al. 2013). Among the 17 species of metallic, pollen-collecting Dialictus (sensu Gibbs 2013; see also Gibbs 2010, 2011), the ground-nesting species L. nymphaearum (= oceanicum (Cockerell); see Gibbs 2010, 2011], L. zephyrum, and true L. versatum (sensu Gibbs 2010, 2011) were most numerous, followed by the wood-nesting species L. cressonii. Interesting records for the subgenus include L. marinum, a species strongly associated with Atlantic coastal sand dunes, L. nigroviride, a relatively large species with a northern distribution, and four male paratypes of L. rozeni (Gibbs 2011), a presumed social parasite of other Dialictus species. Additional L. (Dialictus) recorded from the region including other Atlantic coastal islands of Northeastern Naturalist 56 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Massachusetts (G.I. Stage, P.Z. Goldstein, and J.S. Ascher, unpubl. data) were not found on GI (e.g., L. pruinosum, L. timothyi, and L. vierecki) or were remarkably scarce (e.g., L. leucocomum and L. pilosum). The additional three species, L. foxii, L. birkmanni (=Evylaeus macoupinensis sensu Mitchell [1960], not L. macoupinense sensu Gibbs et al. [2013]), and L. pectorale, have been classified as Evylaeus (Mitchell 1960) or as black, non-metallic Lasioglossum (Dialictus) (Michener 2007), but are now included in an expanded L. (Hemihalictus) (Gibbs et al. 2013). Andrenidae. The 33 Andrena species collected include many that are widely distributed and numerous along the Atlantic coastal plain. Two pollen specialists (oligoleges) of blueberry (Vaccinium) with elongate faces, A. bradleyi and A. carolina, were found only in the 1976–1977 collections. Seven Andrena specialists of goldenrods and other Asteraceae were found: A. canadensis, A. hirticincta, and A. runcinatae (= robervalensis Mitchell; see Gusenleitner et al. 2005) in subgenus Cnemidandrena (but not the regionally numerous A. nubecula); and A. asteris, A. braccata, A. placata, and A. simplex in subgenus Callandrena sensu lato. The Figure 4. Images of Sphecodes johnsonii male (newly recognized): a. dorsal habitus, b. head in front view; and female: c. dorsal habitus, and d. head in front view. Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 57 Andrena sample from GI may be incomplete, as an additional 39 species known from southern NY (sensu Giles and Ascher 2006) were not recorded, and it is difficult for a general collector to sample the host plants of all vernal species during their brief flight periods. However, the Andrena fauna of GI may also be genuinely impoverished to some degree due to reduced diversity and abundance of herbaceous flowers in the forest understory (see above and Hehre 1977). Perdita (Perdita) bradleyi is a new state record for NY based on a single female collected on 15 Jun 1986. This species has also been collected recently at sandy sites in Connecticut (D.L. Wagner, University of Connecticut, Storrs, CT, and J.S. Ascher, unpubl. data), extending its range to the north. Two additional panurgine bees associated with composites were found, Pseudopanurgus compositarum and Perdita octomaculata, as was Calliopsis andreniformis, which usually visits legumes. Megachilidae. The 20 native megachilid species found included one female of the cleptoparasitic Stelis (Dolichostelis) louisae and two males of its host Megachile (Chelostomoides) campanulae (Parker et al. 1987; a report therein of M. angelarum Cockerell as a host of S. louisae in Alabama is implausible as the putative host species is restricted to western North America; see Hurd 1979). Four Osmia species were found, all native, of which two have restricted ranges in NY. The 1976–1977 collections included O. simillima, while the 2006 and 2009 samples included O. chalybea. At this latitude, O. chalybea, a large species associated with Cirsium, is scarce and restricted to the immediate coast, but the species is more common in Florida (where it has a cleptoparasite, Stelis ater Mitchell; Rozen and Hall 2011) and elsewhere on the coastal plain of the southeastern US west to central Texas. Among Megachilini, co-occurrence of Megachile (Leptorachis) petulans and Coelioxys (Glyptocoelioxys) germana is noteworthy, as both of these species and their respective subgenera are at (C. germana) or near (C. petulans) the northeastern limits of their known ranges. A host-parasite association seems very likely and is supported by matching flight seasons (Fig. 3). Two species of subgenus Litomegachile, M. brevis and M. mendica, were abundant, as was Coelioxys sayi, a presumed cleptoparasite of one or both of these Litomegachile at this site, but the larger, consubgeneric M. texana Cresson was not found despite its great abundance in urban New York City (Matteson et al. 2008). Although megachilids comprise a large proportion of exotic bee species in New York State (Matteson et al. 2008), Megachile sculpturalis was the only one found on GI. Apidae. The large carpenter bee Xylocopa virginica was consistently present around buildings on the island, but relatively few were collected. The small carpenter bees Ceratina calcarata and C. dupla (sensu Rehan and Sheffield 2011) were also numerous, but the smaller C. strenua Smith was not collected despite being common elsewhere in southern NY. The 15 species of Nomada recorded included two potentially undescribed morphospecies. Nomada articulata in the erigeronis species group is a cleptoparasite of Agapostemon (Eickwort and Abrams 1980). Of the three autumnal parasites of composite specialist Andrena, Nomada vicina is very likely a parasite of A. hirticincta Northeastern Naturalist 58 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 based in part on matching flight seasons (Fig. 3; a published association with Andrena vicina by Packard 1878, must be erroneous as the flight season of the putative host and its parasite do not overlap). Nomada banksi is possibly a parasite of A. asteris; whereas N. electa is very likely a parasite of A. braccata based on co-occurrence and matching flight seasons at GI (Fig. 3) and at many sites in coastal New England including offshore islands. These potential host associations require confirmation by rearing of parasites from host nests. At least nine additional identified Nomada species and two additional morphospecies in the ruficornis species group are known or presumed to be vernal cleptoparasites of Andrena. The exact number and identity of these remain somewhat uncertain pending further integrative taxonomic studies of this group using DNA markers together with morphological characters such as the arrangement of tibial spines (Droege et al. 2010). A female collected on 28 April 2006 resembles N. composita Mitchell but has a different pseudopygidial fimbria. This morphospecies also occurs on Martha’s Vineyard (P.Z. Goldstein and J.S. Ascher, unpubl. data) and is likely undescribed. Two males of another potentially undescribed species with a subapical mandibular tooth resemble N. maculata in color pattern but are smaller and have single bladelike spines at the apex of each hind tibia rather than opposing prong-like spines. Another related morphospecies represented by a single female is tentatively identified as Nomada cf. perplexa Cresson. The cleptoparasite Triepeolus pectoralis was collected on goldenrod along with its host Melissodes druriella. The larger species Melissodes desponsa was collected in large numbers (213 specimens) on its Cirsium (thistle) host plants, but its cleptoparasite Triepeolus donatus was not found. Four Epeolus cleptoparasites of Colletes included Epeolus lectoides, a species rarely recorded in NY, where it is known exclusively from coastal sites within the range of its presumed host C. nudus. It was first recorded from GI by Bequaert in August 1918 (Leonard 1928). Epeolus scutellaris is likely associated with C. simulans armatus based on frequent co-occurrence in places such as upstate NY where their congeners are scarce. Based on size and season of flight, it would seem that the larger and lateflying E. autumnalis should be associated with C. compactus compactus, and the smaller E. pusillus should be associated with C. americanus (and potentially its close relative C. speculiferus), although the published association for E. pusillus is with C. compactus compactus (Rozen 1966). A single female squash bee Peponapis pruinosa was obtained in a general collection (not on squash). Habropoda laboriosa, a vernal blueberry pollinator represented by a single male, reaches its northeastern range limit in Long Island (including Brooklyn in New York City) and southern Connecticut. Another species of Anthophorini, Anthophora walshii, was not found on GI but was confirmed for NY based on a specimen in the AMNH collected at nearby Montauk Point on 8 August 1937. The few other records of this species from the northeastern United States come from coastal areas and islands of Massachusetts and Rhode Island. Presence of the Hibiscus specialist Ptilothrix bombiformis is explained by abundance of its host plant along pond margins. This species is near Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 59 its northern range limits on GI, and has also been found in New England (new record) based on three females collected by C.L. Remington on 22 Aug 1996 from West River along Marginal Drive, New Haven County, CT and deposited in the Yale Peabody collection. Three bumble bee species recorded from GI by Leonard (1928) based on records from August 1918 by J. Bequaert included Bombus (Thoracobombus) pensylvanicus (cited as B. americanorum), now scarce in the state and region (Giles and Ascher 2006) but still numerous in southernmost New Jersey (Cape May County), in addition to B. (Pyrobombus) vagans and the ubiquitous B. impatiens. Bombus pensylvanicus is absent in recent collections from GI but is known from a specimen in the Museum of Comparative Zoology (examined by J.S. Ascher) from 1918 and a worker in the AMNH collected on 11 Jun 1922 by F.M. Schott. Of the eight bumble bee species collected recently on GI, B. (Cullumanobombus) griseocollis was the most abundant, followed closely by B. (Pyrobombus) impatiens and its social parasite B. (Psithyrus) citrinus (also known to attack other Bombus species). A large number of the relatively long-tongued B. (Thoracobombus) fervidus was collected, and this species persists in NYC as well in larger parks and at other sites with clovers. Bombus (Pyrobombus) perplexus was found to be abundant on GI as was Bombus vagans vagans, a relatively scarce and local species in and around NYC. Perhaps the most surprising of all bees collected on GI was a single B. (Cullumanobombus) rufocinctus worker. This species has a limited distribution in NY, being locally numerous in the Albany area and also expected at certain northern sites but not along the Atlantic coast (Leonard 1928). It is known to occur locally on Cape Cod, MA in cranberry bogs (MacKenzie and Averill 1995). Ecological, Behavioral, and Taxonomic Patterns Ecological and behavioral patterns for both bee individuals and species as documented in the GI dataset (Fig. 5) are generally consistent with those reported in previous studies from the region (Giles and Ascher 2006, Ginsberg 1983), such as large numbers of eusocial individuals relative to eusocial species, larger numbers of parasitic species relative to parasitic individuals, high abundance and diversity of Halictinae and Andreninae (Andrena), and a preponderance of soil-nesting polylectic species and individuals. Among pollen-specialist species collected were at least 17 oligoleges or mesoleges (Cane and Sipes 2006) of Asteraceae including 5 Colletes, 7 Andrena, 2 Panurginae, and 3 Melissodes. These composite-specialist bees collectively host an impressive diversity of Nomadine cleptoparasites, including 3 Nomada and all five Epeolini. Thus, no fewer than 25 bee species (17% of all species) found on Figure 5 (following page). All identified specimens excluding Apis mellifera and Bombus pensylvanicus sorted by: a. nesting substrate by individuals, b. nesting substrate by species, c. sociality by individuals, d. sociality by species, e. level of host plant specialization by individuals, f. level of host plant specialization by species, g. subfamily by individuals, and h. subfamily by species. To accommodate text, the smallest sections of g and h are not to scale. Northeastern Naturalist 60 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 61 GI depend on summer- or fall-blooming Asteraceae (especially Solidago sensu lato, despite records for only four species in the flora) and have correspondingly late and restricted flight seasons (Fig. 2). In addition to these specialists, many polylectic species, including Halictinae, Megachilini, and Osmia chalybea, were most frequently collected on composites. The Vaccinium specialists Andrena carolina and A. bradleyi were represented only by a few specimens from the 1970s. Several species that nest in sandy substrates were present (Colletes spp., Lasioglossum marinum, Agapostemon splendens), but certain sand-associated Lasioglossum (Dialictus) found commonly elsewhere in the region, such as L. vierecki (Crawford), were not recorded. Spring-flying Andrena associated with rosaceous trees and shrubs and other woody plants such as Rhus were well represented, but many species requiring herbaceous forest understory flowers were scarce or absent (cf. Giles and Ascher 2006). Absence of certain oligoletic species found regionally such as specialists on Lyonia ligustrina (L.) DC. can be attributed to an absence of their host plants in the flora (Hehre 1977). Although 18 exotic bee species are recorded from NY, only five were found on GI, including Apis mellifera (European Honey Bee). Hylaeus punctatus was previously known in the northeastern US only from the NYC area, so presence at GI demonstrates spread of this species beyond urban and suburban areas. Lasioglossum (Leuchalictus) leucozonium has long been present in North America but has only recently been recognized as exotic in origin (hypothesized by Giles and Ascher 2006; supported with DNA evidence by Zayed et al. 2007). This species and Andrena wilkella are ground-nesters, whereas Megachile sculpturalis nests in cavities and is known to usurp and occupy active Xylocopa virginica nests in NY (Laport and Minckley 2012). Bee Records from Small NY Islands near GI Records of bees from small islands of Suffolk County, NY, within a 20-mile radius of GI are of interest. These include 45 specimens collected from Great Gull Island by David A. Brody on five dates during 22 May–1 Aug 1976 and 5 collected by J.C. Pallister 14–15 Jun 1949. Collections from this island comprise 17 species—15 shared with Gardiners Island and two additional species (both from 3 Jul 1976), Lasioglossum (Dialictus) albipenne and Melissodes (Eumelissodes) subillata, the latter of which is not known from other northeastern coastal islands. Collections of 35 bee species were made from Fishers Island by Rev. J.L. Zabriskie on ten dates during 1877–1878, 1890–1891, and 1920. The 12 species found are shared with Gardiners Island excepting L. albipenne and Bombus (Pyrobombus) ternarius. The latter record is unexpected as B. ternarius is considered to be a northern or montane species in New York (Leonard 1928) and Connecticut (Viereck 1916) and is otherwise unrecorded from the coast, although it has been found recently as far south as Black Rock Forest, Orange County (Giles and Ascher 2006) and is regularly found at coastal sites south to Boston, MA, and vicinity (S. Kent, Boston, MA, pers. comm.). Northeastern Naturalist 62 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Comparison to the Broader Fauna In comparison to the bee fauna of NYS as a whole (447 described species in 45 genera), and to some well-sampled localities within the state, the 155 species and morphospecies known from GI species are relatively few, but still impressive for a small island sampled using only nets, primarily by a single non-specialist collector. Most species recorded are widely distributed in eastern North America, and are well known from NY, the other mid-Atlantic states, and southern New England. However, several specimens from GI are of considerable interest as they represent new state records or are among the only known recent collections from the state. Several species found at GI are well known from the mid-Atlantic coast south of NY but have been infrequently collected in NY and southern New England. These “southern” or coastal species include Colletes nudus and its presumed parasite Epeolus lectoides, Perdita bradleyi, Osmia chalybea, Megachile petulans and its presumed parasite Coelioxys germana, Ptilothrix bombiformis, and Habropoda laboriosa. The few “northern” species at GI include a disjunct record of Bombus rufocinctus. Gardiners Island has far fewer exotic bee species and individuals than were recorded from suburban (Fetridge et al. 2008) and urban (Matteson et al. 2006) gardens of southern NY, although two exotic species detected recently in the state have already colonized it in addition to three ubiquitous, long-established species. Acknowledgments Jason Gibbs made or confirmed identifications for Lasioglossum (Dialictus) in the course of his revisionary studies. Sandra Rehan identified many female Ceratina calcarata and dupla. Mike Arduser shared unpublished diagnostic information about Sphecodes including S. johnsonii. Maximilian Schwarz and Sam Droege shared unpublished data on the taxonomy of Pseudoanthidium and Nomada, respectively. Hadel Go photographed S. johnsonii. Douglas Yanega alerted us to the Dianthidium simile record. P. Gambino and Kevin Matteson provided unpublished bee records from New York. Gerald I. Stage and Paul Z. Goldstein shared unpublished data on bees of Massachusetts coastal islands, and Tracy Zarillo shared unpublished data on the bees of coastal Connecticut. Databasing of bees from Gardiners Island was made possible by generous support from R.G. Goelet and from NSFDBI# 0956388, Collaborative Research: Collaborative Databasing of North American Bee Collections within a Global Informatics Network, PI J.S. Ascher, Co-PIs Jerome G. Rozen, Jr. and Douglas Yanega. We thank Charles Elias and Margaret A. Rozen for databasing bee specimens from the island. Paul Z. Goldstein, Jerome G. Rozen, Jr., and Eli S. Wyman commented on the manuscript. Literature Cited Aguib, S., K. Louadi, and M. Schwarz. 2010. Les Anthidiini (Megachilidae, Megachilinae) d’Algérie avec trois espèces nouvelles pour ce pays: Anthidium (Anthidium) florentinum (Fabricius, 1775), Anthidium (Proanthidium) amabile Alfken, 1932, et Pseudoanthidium (Exanthidium) enslini (Alfken, 1928). Entomofauna 31(12):121–152. Arthropod Easy Capture. 2013. Version: 1.34. Available online at http://sourceforge.net/p/ arthropodeasy. Accessed 18 February 2013. Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 63 Ascher, J.S., and J. Pickering. 2013. Discover Life bee species guide and world checklist (Hymenoptera: Apoidea: Anthophila). [Draft 35, 20 Mar 2013]. Available online at http://www.discoverlife.org/mp/20q?guide=Apoidea_species&flags=HAS. Ascher, J.S., P. Gambino, and S. Droege. 2006. Adventive Hylaeus (Spatulariella Popov) in the New World (Hymenoptera: Apoidea: Colletidae). Proceedings of the Entomological Society of Washington 108(1):237–239. 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Floral specialization by bees: Analytical methods and a revised lexicon for oligolecty. Pp. 99–122, In N.M. Waser and J. Ollerton (Eds.). Plant- Pollinator Interactions: From Specialization to Generalization. University of Chicago Press, Chicago, IL. Colla, S.R., J.S. Ascher, M. Arduser, J. Cane, M. Deyrup, S. Droege, J. Gibbs, T. Griswold, H. G. Hall, J. Neff, R.P. Jean, M.G. Rightmyer, C. Sheffield, M. Veit, and A. Wolf. 2012. Documenting persistence of most eastern North American bee species (Hymenoptera: Apoidea: Anthophila) to 1990–2009. Journal of the Kansas Entomological Society 85(1):14–22. Droege, S.W., and L.H. Shapiro. 2011. An August survey of wild bees (Hymenoptera: Apoidea) in the northeastern port areas of Baltimore, Maryland and the second North American Record of Pseudoanthidium nanum (Mocsáry). Maryland Entomologist 5(3):33–44. Droege, S., M.G. Rightmyer, C.S. Sheffield, and S.G. Brady. 2010. New synonymies in the bee genus Nomada from North America (Hymenoptera: Apidae). Zootaxa 2661:1–32. Eickwort, G.C., and J. Abrams. 1980. Parasitism of sweat bees in the genus Agapostemon by cuckoo bees in the genus Nomada (Hymenoptera: Halictidae, Anthophoridae). Pan- Pacific Entomologist 56:144–152. Fetridge, E.D., J.S. Ascher, and G.A. Langellotto. 2008. The bee fauna of residential gardens in a suburb of New York City (Hymenoptera Apoidea). Annals of the Entomological Society of America 101(6):1067–1077. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus) in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa 2591:1–382. Gibbs, J. 2011. Revision of the metallic Lasioglossum (Dialictus) of eastern North America (Hymenoptera: Halictidae: Halictini). Zootaxa 3073:1–216. Gibbs, J.S., L. Packer, S. Dumesh, and B.N. Danforth. 2013. Revision and reclassification of Lasioglossum (Evylaeus), L. (Hemihalictus), and L. (Sphecodogastra) in eastern North America (Hymenoptera: Apoidea: Halictidae). Zootaxa 3672:1–117. Northeastern Naturalist 64 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Giles, V., and J.S. Ascher. 2006. Bees of the Black Rock Forest Preserve, New York (Hymenoptera: Apoidea). Journal of Hymenoptera Research 15(2):208–231. Ginsberg, H.S. 1983. Foraging ecology of bees in an old field. Ecology 64:165–175. Grixti, J.C., L.T. Wong, S.A. Cameron, and C. Favret. 2008. Decline of bumble bees (Bombus) in the North American Midwest. Biological Conservation 142(1):75–84. Gusenleitner, F., M. Schwarz, J.S. Ascher, and E. Scheuchl. 2005. Korrekturen und Nachträge zu Gusenleitner & Schwarz (2002): Weltweite Checkliste der Bienengattung Andrena mit Bemerkungen und Ergänzungen zu paläarktischen Arten (Hymenoptera: Apidae, Andreninae, Andrena) Entomofauna 26(26):437–472. Hehre, E J. 1977. The flora of Gardiners Island. Rhodora 79:214–239. Hurd, P.D., Jr. 1979. Superfamily Apoidea. 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Hewson-Smith, and J.S. Ascher. 2010. Specimen databases: A case study in entomology using web-based software. American Entomologist 56(4):206–216. Sheffield, C.S., S. Dumesh, and M. Cheryomina. 2011. Hylaeus punctatus (Hymenoptera: Colletidae), a bee species new to Canada with a review of other non-native species. Journal of the Entomological Society of Ontario 142:29–43. Viereck, H.L. 1916. Apoidea. The Hymenoptera or, wasp-like insects, of Connecticut. State of Connecticut Geological and Natural History Survey Bulletin No. 22:1–824 + 10 Pls, 15 figs. Zayed, A., S.A. Constantin, and L. Packer. 2007. Successful biological invasion despite a severe genetic load. PLoS ONE 2:e868. Northeastern Naturalist 66 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Appendix 1. Species list of bees recorded from Gardiners Island (GI) and neighboring Fishers Island (FI), and Great Gull Island (GG) including totals of databased specimens for New York State (NY), New Jersey (NJ), and Connecticut (CT) for comparison. Numbers in brackets pertain to specimens that were not recorded in the database. For “nest” substrate: C = cavity, H = hive, P = pithy stems, S = soil. The following superscripted annotations clarify aspects of nest type: c = cells of cellophane-like material, r = lined with resin, l = lined with leaves or other plant materials, w = solid wood (all other wood nesters excavate rotten wood). Degree of sociality: B = subsocial, E = eusocial (all primitively eusocial except the advanced eusocial Apis mellifera; some Halictus and other halictines are known to have solitary populations across their ranges), P = parasitic (all cleptoparasitic except the Lasioglossum and Bombus which are known or suspected to be socially parasitic; the status of most Sphecodes is unclear), and S = solitary (and/or communal in the case of Agapostemon or Lasioglossum subgenus Lasioglossum). Status: O = oligolectic, P = parasitic, E = exotic. Family/species GI NY NJ CT FI GG Nest Sociality Status Family Colletidae Colletes americanus Cresson, 1868 96 162 17 28 0 0 S S Colletes compactus compactus Cresson, 1868 220 377 18 54 0 0 S S O Colletes inaequalis Say, 1837 5 489 194 115 0 0 S S Colletes nudus Robertson, 1898 54 54 0 0 0 0 S S Colletes simulans armatus Patton, 1879 231 385 37 45 0 0 S S O Colletes solidaginis Swenk, 1906 1 2 2 8 0 0 S S Colletes speculiferus Cockerell, 1927 [= C. mitchelli Stephen] 7 15 5 1 0 0 S S Colletes thoracicus Smith, 1853 259 329 141 67 0 0 S S Hylaeus (Hylaeus) mesillae cressoni (Cockerell, 1907) 5 290 97 141 1 3 Cc S Hylaeus (Prosopis) affinis (Smith, 1853) 4 85 19 56 1 0 Cc S Hylaeus (Prosopis) modestus modestus Say, 1837 65 501 89 183 1 0 Cc S Hylaeus (Prosopis) cf. nelumbonis (Robertson, 1890) 2 3 0 0 0 0 Cc S Hylaeus (Spatulariella) punctatus (Brullé, 1832) [1] 33 46 0 0 0 Cc S E Family Halictidae Augochloropsis (Paraugochloropsis) metallica (F., 1793) 84 198 32 61 0 0 S S Augochlorella aurata (Smith, 1853) 533 2752 223 1322 8 4 S E Augochlora (Augochlora) pura pura (Say, 1837) 78 1078 171 273 0 0 W S Agapostemon (Agapostemon) sericeus (Förster, 1771) 157 446 63 127 1 0 S S Northeastern Naturalist Vol. 21, No. 1 J.S. 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Goelet 2014 67 Family/species GI NY NJ CT FI GG Nest Sociality Status Agapostemon (Agapostemon) splendens (Lepeletier, 1841) 8 50 6 24 1 0 S S Agapostemon (Agapostemon) texanus Cresson, 1872 3 40 4 129 0 0 S S Agapostemon (Agapostemon) virescens (Fabricius, 1775) 519 1254 134 571 2 4 S S Sphecodes aroniae Mitchell, 1960 7 20 14 5 0 0 S P P Sphecodes confertus Say, 1837 1 63 18 50 0 0 S P P Sphecodes coronus Mitchell, 1956 4 116 2 22 0 0 S P P Sphecodes davisii Robertson, 1897 4 67 9 36 0 0 S P P Sphecodes dichrous Smith, 1853 4 37 6 43 0 0 S P P Sphecodes heraclei Robertson, 1897 47 58 6 11 0 0 S P P Sphecodes johnsonii Lovell, 1909 21 45 1 20 0 0 S P P Sphecodes mandibularis Cresson, 1872 8 30 2 10 0 0 S P P Sphecodes pimpinellae Robertson, 1900 4 16 0 5 0 0 S P P Sphecodes ranunculi Robertson, 1897 1 53 2 16 0 0 S P P Halictus (Nealictus) parallelus Say, 1837 30 33 1 30 0 0 S E Halictus (Odontalictus) ligatus Say, 1837 977 2166 458 1359 4 2 S E Halictus (Protohalictus) rubicundus (Christ, 1791) 17 526 43 126 0 0 S E Halictus (Seladonia) confusus confusus Smith, 1853 123 1428 310 437 0 1 S E [Lasioglossum (Dialictus) albipenne (Robertson, 1890)] 0 10 4 3 1 5 W* E Lasioglossum (Dialictus) bruneri (Crawford, 1902) 13 133 20 57 0 0 S E Lasioglossum (Dialictus) coeruleum (Robertson, 1893) 1 200 5 37 0 0 W E Lasioglossum (Dialictus) cressonii (Robertson, 1890) 80 491 49 267 0 4 W E Lasioglossum (Dialictus) imitatum (Smith, 1853) 6 741 616 216 0 0 S E Lasioglossum (Dialictus) laevissimum (Smith, 1853) 0 73 0 5 0 0 S E Lasioglossum (Dialictus) leucocomum (Lovell, 1908) 1 78 72 197 0 0 S E Lasioglossum (Dialictus) lineatulum (Crawford, 1906) 1 1001 63 21 0 0 S E Lasioglossum (Dialictus) marinum (Crawford, 1904) 6 65 3 61 0 0 S E Lasioglossum (Dialictus) nigroviride (Graenicher, 1911) 3 84 3 23 0 0 S E Lasioglossum (Dialictus) nymphaearum (Robertson, 1895) 364 441 84 109 3 3 S E Lasioglossum (Dialictus) oblongum (Lovell, 1905) 5 101 3 135 0 0 W E Lasioglossum (Dialictus) pilosum (Smith, 1853) 4 176 248 689 8 0 S E Northeastern Naturalist 68 J.S. 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Goelet 2014 Vol. 21, No. 1 Family/species GI NY NJ CT FI GG Nest Sociality Status Lasioglossum (Dialictus) rozeni Gibbs, 2011 13 18 0 0 0 0 S P P Lasioglossum (Dialictus) subviridatum (Cockerell, 1938)** 2 299 3 4 0 0 W E Lasioglossum (Dialictus) tegulare (Robertson, 1890) 18 343 294 345 0 0 S E Lasioglossum (Dialictus) versatum (Robertson, 1902) 1752 2195 26 714 0 1 S E Lasioglossum (Dialictus) zephyrum (Smith, 1853) 172 449 82 47 0 0 S E Lasioglossum (Hemihalictus) birkmanni (Crawford, 1906) 1 102 10 12 0 0 S S Lasioglossum (Hemihalictus) foxii (Robertson, 1895) 1 824 17 96 0 0 S S Lasioglossum (Hemihalictus) pectorale (Smith, 1853) 93 210 40 247 0 0 S S Lasioglossum (Lasioglossum) coriaceum (Smith, 1853) 55 185 20 287 0 3 S S Lasioglossum (Lasioglossum) fuscipenne (Smith, 1853) 33 71 25 41 0 0 S S Lasioglossum (Leuchalictus) leucozonium (Schrank, 1781) 8 157 12 72 0 0 S S E Lasioglossum (Sphecodogastra) quebecense (Crawford, 1907) 7 533 13 62 0 0 S S Family Andrenidae Andrena (Andrena) carolina Viereck, 1909 3 221 5 68 0 0 S S O Andrena (Andrena) frigida Smith, 1853 5 110 9 54 0 0 S S O Andrena (Andrena) mandibularis Robertson, 1892 21 75 5 11 0 0 S S Andrena (Andrena) tridens Robertson, 1902 33 53 0 18 0 0 S S Andrena (Callandrena s.l.) asteris Robertson, 1891 59 119 2 53 0 0 S S O Andrena (Callandrena s.l.) braccata Viereck, 1907 105 134 23 28 0 0 S S O Andrena (Callandrena s.l.) placata Mitchell, 1960 1 72 16 36 0 0 S S O Andrena (Callandrena s.l.) simplex Smith, 1853 1 96 14 24 0 0 S S O Andrena (Cnemidandrena) canadensis Dalla Torre, 1896 13 56 7 18 0 0 S S O Andrena (Cnemidandrena) hirticincta Provancher, 1888 107 406 29 132 0 0 S S O Andrena (Cnemidandrena) runcinatae Cockerell, 1906 2 18 0 1 0 0 S S O Andrena (Conandrena) bradleyi Viereck, 1907 2 117 54 61 0 0 S S O Andrena (Holandrena) cressonii cressonii Robertson, 1891 33 345 33 67 0 0 S S Andrena (Larandrena) miserabilis Cresson, 1872 90 1457 120 233 0 0 S S Andrena (Leucandrena) barbilabris (Kirby, 1802) [1] 174 28 59 0 0 S S Andrena (Melandrena) carlini Cockerell, 1901 23 1075 82 406 0 0 S S Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 69 Family/species GI NY NJ CT FI GG Nest Sociality Status Andrena (Melandrena) commoda Smith, 1879 2 86 7 6 0 0 S S Andrena (Melandrena) pruni Robertson, 1891 15 141 15 61 0 0 S S Andrena (Melandrena) vicina Smith, 1853 42 717 83 338 0 0 S S Andrena (Micrandrena) neonana Viereck, 1917 6 7 0 2 0 0 S S Andrena (Plastandrena) crataegi Robertson, 1893 43 2411 102 584 0 3 S S Andrena (Rhacandrena) brevipalpis Cockerell, 1930 13 166 14 22 0 0 S S Andrena (Scrapteropsis) alleghaniensis Viereck, 1907 9 157 41 32 0 0 S S Andrena (Scrapteropsis) ilicis Mitchell, 1960 2 3 3 0 0 0 S S Andrena (Scrapteropsis) imitatrix Cresson, 1872 149 624 36 99 0 0 S S Andrena (Simandrena) nasonii Robertson, 1895 170 1744 64 320 0 0 S S Andrena (Taeniandrena) wilkella (Kirby, 1802) 26 2822 131 232 0 1 S S E Andrena (Trachandrena) forbesii Robertson, 1891 51 656 28 63 0 0 S S Andrena (Trachandrena) hippotes Robertson, 1895 196 763 29 83 0 0 S S Andrena (Trachandrena) nuda Robertson, 1891 95 159 15 32 0 0 S S Andrena (Trachandrena) rugosa Robertson, 1891 2 494 10 117 0 0 S S Andrena (Trachandrena) spiraeana Robertson, 1895 4 151 5 26 0 0 S S Andrena (Tylandrena) perplexa Smith, 1853 29 269 14 34 0 0 S S Calliopsis (Calliopsis) andreniformis Smith, 1853 2 181 147 197 0 0 S S Pseudopanurgus compositarum (Robertson, 1893) 3 10 5 1 0 0 S S O Perdita (Alloperdita) bradleyi Viereck, 1907 1 1 1 13 0 0 S S Perdita (Perdita) octomaculata (Say, 1824) 1 216 13 42 0 0 S S O Family Megachilidae Stelis (Dolichostelis) louisae Cockerell, 1911 1 7 3 1 0 0 C P Hoplitis (Alcidamea) pilosifrons (Cresson, 1864) 13 32 15 40 0 0 P S Hoplitis (Alcidamea) producta producta (Cresson, 1864) 1 147 20 63 0 0 P S Osmia (Helicosmia) chalybea Smith, 1853 8 8 2 0 0 0 C S Osmia (Melanosmia) atriventris Cresson, 1864 2 116 19 36 0 0 C S Osmia (Melanosmia) pumila Cresson, 1864 19 310 51 195 0 0 C S Osmia (Melanosmia) simillima Smith, 1853 5 14 0 3 0 1 C S Northeastern Naturalist 70 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 Vol. 21, No. 1 Family/species GI NY NJ CT FI GG Nest Sociality Status Megachile (Callomegachile) sculpturalis Smith, 1853 5 144 2 7 0 0 Cr S E Megachile (Chelostomoides) campanulae (Robertson, 1903) 2 104 8 75 0 0 Cr S Megachile (Leptorachis) petulans Cresson, 1878 151 157 8 0 0 0 Cl S Megachile (Litomegachile) brevis Say, 1837 38 77 17 87 1 1 Cl S Megachile (Litomegachile) mendica Cresson, 1878 125 433 118 107 0 0 Cl S Megachile (Megachile) montivaga Cresson, 1878 1 3 5 17 0 0 Cl S Megachile (Sayapis) inimica sayi Cresson, 1878 1 25 4 3 0 0 Cl S Megachile (Xanthosarus) frigida Smith, 1853 1 63 2 6 0 0 Cl S Megachile (Xanthosarus) gemula gemula Cresson, 1878 3 113 15 36 0 0 Cl S Megachile (Xanthosarus) latimanus Say, 1823 14 301 50 41 2 0 Cl S Coelioxys (Boreocoelioxys) octodentata Say, 1824 1 49 6 11 0 0 C P P Coelioxys (Boreocoelioxys) rufitarsis Smith, 1854 1 77 2 15 0 0 C P P Coelioxys (Boreocoelioxys) sayi Robertson, 1897 115 186 26 17 0 0 C P P Coelioxys (Glyptocoelioxys) germana Cresson, 1878 40 42 4 0 0 0 C P P Family Apidae Xylocopa (Xylocopoides) v. virginica (Linnaeus, 1771) 132 476 110 141 0 0 Ww B Ceratina (Zadontomerus) calcarata Robertson, 1900 167 695 178 469 0 6 P B Ceratina (Zadontomerus) dupla Say, 1837 82 240 52 360 1 4 P B Nomada articulata Smith, 1854 26 119 29 65 0 0 S P P Nomada banksi Cockerell, 1907 1 5 1 0 0 0 S P P Nomada sp. nr. composita Mitchell, 1962 1 1 0 0 0 0 S P P Nomada cressonii Robertson, 1893 8 209 27 105 0 1 S P P Nomada denticulata Robertson, 1902 3 58 18 11 0 0 S P P Nomada electa Cresson, 1863 48 56 2 0 0 0 S P P Nomada illinoensis Robertson, 1900 2 62 3 19 0 0 S P P Nomada imbricata Smith, 1854 7 82 5 13 0 0 S P P Nomada maculata Cresson, 1863 1 149 70 178 0 0 S P P Nomada parva Robertson, 1900 12 14 6 4 0 0 S P P Nomada cf. perplexa Cresson, 1863 1 18 9 6 0 0 S P P Northeastern Naturalist Vol. 21, No. 1 J.S. Ascher, S. Kornbluth, and R.G. Goelet 2014 71 Family/species GI NY NJ CT FI GG Nest Sociality Status Nomada pygmaea Cresson, 1863 13 144 26 44 0 0 S P P Nomada sayi Robertson, 1893 1 18 8 5 0 0 S P P Nomada vicina Cresson, 1863 23 39 3 11 0 0 S P P Nomada sp. [with blade-like hind tibial setae] 1 13 0 0 0 0 S P P Epeolus autumnalis Robertson, 1902 6 13 4 2 0 0 S P P Epeolus lectoides Robertson, 1901 24 25 1 1 0 0 S P P Epeolus pusillus Cresson, 1864 14 29 8 6 0 0 S P P Epeolus scutellaris Say, 1824 22 63 17 5 0 0 S P P Triepeolus pectoralis (Robertson, 1897) 5 47 9 1 0 0 S P P Ptilothrix bombiformis (Cresson, 1878) 2 22 14 0 0 0 S S O Melissodes (Eumelissodes) druriella (Kirby, 1802) 49 161 25 31 0 0 S S O [Melissodes (Eumelissodes) subillata LaBerge, 1961] 0 33 10 10 0 1 S S O Melissodes (Eumelissodes) trinodis Robertson, 1901 2 26 14 15 0 0 S S O Melissodes (Heliomelissodes) desponsa Smith, 1854 196 243 10 14 1 0 S S O Peponapis (Peponapis) pruinosa (Say, 1837) 1 187 110 64 2 0 S S O Habropoda laboriosa (Fabricius, 1804) 1 13 27 3 0 0 S S O Anthophora (Clisodon) terminalis Cresson, 1869 1 89 18 36 0 0 W S Bombus (Psithyrus) citrinus (Smith, 1854) 154 546 25 87 0 0 H P P Bombus (Thoracobombus) fervidus (Fabricius, 1798) 78 526 76 160 0 0 H E Bombus (Thoracobombus) pensylvanicus (DeGeer, 1773) 1 141 66 68 0 0 H E Bombus (Cullumanobombus) rufocinctus Cresson, 1863 1 15 0 0 0 0 H E Bombus (Cullumanobombus) griseocollis (DeGeer, 1773) 587 869 139 105 0 3 H E Bombus (Pyrobombus) bimaculatus Cresson, 1863 133 371 227 295 0 0 H E Bombus (Pyrobombus) impatiens Cresson, 1863 459 1754 763 1295 0 0 H E Bombus (Pyrobombus) perplexus Cresson, 1863 29 627 129 279 0 0 H E [Bombus (Pyrobombus) ternarius Say, 1837] 0 367 1 13 1 0 H E Bombus (Pyrobombus) vagans vagans Smith, 1854 81 752 111 302 0 0 H E Apis (Apis) mellifera Linnaeus, 1758 1 61 9 679 0 0 H E E *Wood-nesting by L. albipenne is uncertain but consistent with behavior of the similar L. cressonii. **Most specimens reported as L. oblongum in Giles and Ascher (2006) pertain to this species, the status of which was clarified by Gibbs (2011) who noted its wood-nesting behavior.