The Bee Fauna of Inland Sand Dune and Ridge Woodland
Communities in Worcester County, Maryland
Jennifer A. Selfridge, Christopher T. Frye, Jason Gibbs, and Robert P. Jean
Northeastern Naturalist, Volume 24, Issue 4 (2017): 421–445
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2017 NORTHEASTERN NATURALIST 24(4):421–445
The Bee Fauna of Inland Sand Dune and Ridge Woodland
Communities in Worcester County, Maryland
Jennifer A. Selfridge1,*, Christopher T. Frye1, Jason Gibbs2, and Robert P. Jean3
Abstract - We surveyed bees inhabiting inland dune and ridge woodlands at 30 sites in
Worcester County, MD, in 2008 and 2009 . We collected and identified 4878 bees representing
5 families, 31 genera, and 121 species. Here, we report data on annual and seasonal
variation. Expanding survey efforts to include multiple years and seasons served to increase
the overall number of species encountered, primarily through documenting the presence
of rare or infrequently collected species. Eighty-eight species (73%) were represented by
fewer than 10 individuals; of these 30 were represented by a single individual (25% of the
total). The 5 most dominant species represented nearly half (48%) of the total number of
specimens. We report a list of bee species collected from inland dune and ridge woodlands
and discuss the presence of apparently habitat-restricted species.
Introduction
Xeric habitats are characterized by dry conditions with persistently low moisture
levels and they often support specialist and habitat-restricted invertebrate
species that are uniquely adapted to such conditions (Cloudsley-Thompson 1975).
Reasons for habitat restriction may include specialization on limited-range host
plants (Litvaitis et al. 1999, Wagner et al. 2003); unique microhabitat availability
for nesting, burrowing, and foraging (Droege et al. 2009, Litvaitis et al. 1999,
Wagner et. al 2003); and avoidance of predators and parasitoids (Fernandes and
Price 1992). Frye et al. (2014) discussed butterfly, beetle, and leafhopper species
restricted to xeric habitats in some detail and provided specific examples pertaining
to Maryland species. Inland dune and ridge woodlands (Fig. 1), herein referred
to generally as “dunes” or “dune habitat”, are globally rare xeric habitats that
occur only on the Delmarva Peninsula and in southern New Jersey (NatureServe
2017). These communities are characterized by low-relief inland dunes shaped
by northwest winds during the Pleistocene epoch and comprised of sand sheets of
the Parsonsburg Formation (Denny et al. 1979, Newell and Dejong 2011). These
woodlands are dominated by Pinus spp. (pines) and Quercus spp. (oaks) and are
described at length by Frye et al. (2014).
The bee fauna of these dune habitats has not been thoroughly investigated.
Although roughly 430 species of native bees have been documented in Maryland
1Maryland Department of Natural Resources, Wildlife and Heritage Service, Natural Heritage
Program, 909 Wye Mills Road, Wye Mills, MD 21601. 2University of Manitoba, Department
of Entomology, Wallis Roughley Museum of Entomology, 214 Animal Science/
Entomology Building, 12 Dafoe Road, Winnipeg, MB R3T 2N2, Canada. 3Environmental
Solutions & Innovations, Inc., 1811 Executive Drive, Suites C–D, Indianapolis, IN 46241.
*Corresponding author - jennifer.selfridge@maryland.gov.
Manuscript Editor: Howard Ginsburg
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to date (Ascher and Pickering 2017), our understanding of the ecological requirements
of most of the species is limited. This knowledge gap is important amidst
concerns of recent pollinator declines (NRCNA 2007) and the listing of 8 bee species
under the Endangered Species Act including 7 Hawaiian Hylaeus species and
Bombus affinis Cresson (Rusty Patched Bumblebee). Many other insects, among
them butterflies, moths, wasps, flies, and beetles, are noted for their value as pollinators
(Kevan and Baker 1983, Rader et al. 2016); however, bees are arguably the
most important insect pollinators because they actively collect pollen. Native bees
play major roles in the pollination of crops (Greenleaf and Kremen 2006a, 2006b;
Winfree et al. 2008) and native plants (Kevan and Baker 1983, Ollerton et al. 2011).
Native bees have received more attention in recent years for their value as pollinators
due to concerns over managed Apis mellifera L. (Western Honey Bee) declines
(Garibaldi et al. 2013, Winfree et al. 2008). Evidence of decline for other bees is
growing (Bartomeus et al. 2013, Biesmeijer et al. 2006, Kremen et al. 2002, Potts
et al. 2010), most notably for Bombus spp. (bumblebees; Cameron et al. 2011, Colla
and Packer 2008, Grixti et al. 2009) because the escape of commercially produced
bumblebees used for greenhouse pollination is thought to be a factor in the spread
of pathogens to wild bees (Colla et al. 2006).
Many bee species prefer to nest in loose, sandy soil (Cane 1991, Michener
2007); thus, dune habitats may be beneficial to many ground-nesting bees. This
Figure 1. A typical example of an inland sand and ridge woodland site (sand-dune habitat)
in Worcester County, MD.
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study documents the bee species present in this rare community type and identifies
those that may be restricted to dune habitats.
Field-site Description
We conducted the study in Worcester County, MD, on the Atlantic Coastal Plain
east of the Chesapeake Bay (Fig. 2). We chose 30 survey sites distributed over
2 adjacent US Geological Survey (USGS) quadrangles (Snow Hill and Dividing
Creek). These areas, known for their dune fields (Newell and Dejong 2011), are
characterized by an increase in elevation above the surrounding forest matrix, an
elliptical shape, and well-drained soils. Dunes are typically interspersed throughout
a landscape of basin swamps and lowland forests.
To locate dunes, we used a combination of USGS quadrangle (topographic)
maps and 2 ArcMap GIS (geographic information system) data layers: US Department
of Agriculture (USDA) Soil Survey Geographic (SSURGO) data, and LIDAR
(light detection and ranging) imagery. These resources allowed us to determine
the locations of dune sites within the 2 quadrangles. We mapped all dunes (n =
303) as polygons using ArcMap GIS. We employed the natural breaks function in
ArcMap (extension X-Tools Pro) to divide the dunes into 3 statistically determined
size-classes (small, medium, and large). We calculated the actual range of each
size class in ArcMap based on natural groupings inherent in the data, determining
Figure 2. Map of the survey area in Worcester County, MD, covering portions of the Dividing
Creek and Snow Hill USGS Quadrangles. Survey sites (dunes) are shown as darkened
areas.
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break points that “best group similar values and maximize the differences between
classes” (ESRI 2016). Of the 303 dunes identified and mapped, we chose 30 at
random for sampling. We ground-truthed all sites to verify that the polygons represented
dune habitat. Of these, 9 were small dunes (<1.1 ha), 14 were medium dunes
(1.1–4.0 ha) and 7 were large dunes (>4.0 ha). In addition to size, sites differed in
management history, forest-stand age, vegetative composition, and nearest-neighbor
distance (distance to other dunes), which varied from 0.03 km to 17 km.
Methods
We sampled bees along 70-m transects using 15 bee bowls, each separated by 5
m, incorporating methodology used in previous studies (e.g., Droege et al. 2010,
Grundel et al. 2011a, Sharpiro et al. 2014, Westphal et al. 2008). Bee bowls offer a
standardized, repeatable method for assessing bee abundance, but have some bias
associated with them because they are known to be inefficient at capturing certain
species (Cane et al. 2000, Geroff et al. 2014, Grundel et al. 2011a, Joshi et al. 2015,
Leong and Thorp 1999, Toler et al. 2005). Each trap consisted of a 96-ml Solo® Brand
plastic soufflé cup Model P325 (7-cm diameter) filled with a 3% Dawn® dish soap
and water solution. We either left bowls white or painted them fluorescent blue or
fluorescent yellow with Guerra Brand paint. We mixed 61 L of silica flat with 473 ml
of fluorescent blue or fluorescent yellow to obtain the desired colors. These traps have
been used successfully by many other bee researchers in the eastern US (e.g., Droege
2006; Grundel et al. 2010, 2011b); capture rates vary depending on region, time of
year, bee species, and other factors (Geroff et al. 2014, Hicks et al. 2012, Leong and
Thorp 1999). We included 5 each of white, yellow, and blue bowls on each transect.
We set 1 transect on small dunes (15 bowls), 2 transects on medium dunes (30 bowls),
and 3 transects on large dunes (45 bowls). We set 4 transects (60 bowls) on the 2
largest dunes (6.9 ha and 10.7 ha). The placement of transects at each dune differed
in 2008 and 2009, but the number of bowls at each dune remained constant. We left
bowls out for a period of 48 h during each spring, summer, and fall sample to limit
mortality and prevent oversampling of the bee populations.
Sampling dates varied slightly from year to year and between different dunes;
the exact dates were dependent upon weather conditions and available labor. We
obtained spring samples between 30 March and 20 May, summer samples between
7 July and 29 July, and fall samples between 18 September and 30 September. We
sampled twice in the spring, once in the summer, and once in the fall in 2008 and 3
times in the spring, once in the summer, and once in the fall in 2009.
When analyzing annual and seasonal data, we excluded species represented
by fewer than 10 individuals to limit the potential bias of rare or infrequently
occurring species. Thus, our final analyses and results included only 27% of the
species collected.
Missing data
We deployed 885 bee bowls across 30 sites in each 48-h sampling period. Occasionally,
individual bowls were lost, most often because they were trampled or
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knocked over by animals. Most dunes incurred significantly less than a 14% loss; we
retrieved 13 of 15 bowls per transect in any given sampling period. In all instances,
there was no systematic bias and we excluded the missing values from the analysis
(see Scheffer 2002); even undisturbed bowls may fail to collect any specimens.
Data analysis
We used abundance data in all analyses because we considered individuals to be
independent of other bees of the same species in each sample. In addition to documenting
overall bee species richness, we compared richness and abundance between
the 3 seasons (spring, summer, and fall) and different years (2008 and 2009).
We employed multiple-response permutation procedure (MRPP) in PC-ORD
(Version 6.22), a non-parametric analog of analysis of variance, to test the null
hypothesis of no significant differences in bee-species composition between years
and seasons. Details of the method can be found in Mielke and Berry (2001). The
strategy of MRPP is to compare the observed intragroup average distances with the
average distances that would have resulted from all the other possible combinations
of the data under the null hypothesis. The test statistic, usually symbolized with a
lowercase delta (δ), is the average of the observed intragroup distances weighted by
relative group size. The observed delta is compared to the possible deltas resulting
from every permutation of the data. The MRPP reports a test statistic (T) describing
the separation between groups, a measure of effect size (A) describing withingroup
agreement, and a P-value representing the likelihood of finding an equal
or smaller delta than the observed based on all possible partitions of the dataset
using the Pearson Type III distribution of deltas. We used Sorenson distance and a
ranked-distance matrix following the protocols in McCune and Grace (2002). We
used indicator-species analysis (ISA) as employed in PC-ORD as a complement to
MRPP to describe the value of different bee species for indicating trends in annual
or seasonal variation. Indicator values range from 0 (no indication) to 100 (perfect
indication). We evaluated statistical significance of indicator values via a Monte
Carlo method using 1000 randomizations.
Species identification and determination of habitat-restricted species
We identified bees with the assistance of Sam Droege (USGS Native Bee
Laboratory, Patuxent Wildlife Research Refuge, Washington, DC) using species
concepts and keys from published revisions (Bouseman and LaBerge 1979; Coelho
2004; Gibbs 2011; Gibbs et al. 2013; LaBerge 1961, 1967, 1971, 1973, 1986;
McGinley 1986; Mitchell 1960, 1962; Rehan and Sheffield 2011; Ribble 1967;
Schwarz and Gusenleitner 2004; Stephen 1954; Williams et al. 2014) and online
resources (e.g., Droege 2016, Larkin et al. 2016). We verified some species identifications
by comparison with expertly identified material in the National Museum of
Natural History, Smithsonian Institution, Washington, DC. Classification follows
Michener (2007) and Ascher and Pickering (2017), and incorporates recent taxonomic
and nomenclatural changes (e.g., Gibbs et al. 2013, Williams et al. 2008).
Available information is limited for many species; thus, we identified rare and/or
habitat-restricted species based on collection data from across the state and region
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using information collected and/or compiled by the US Geological Survey Native
Bee Inventory and Monitoring Lab and Ascher and Pickering (2017).
Results
We collected and identified 4878 bees representing 5 families, 31 genera, and
121 species (Appendix 1). We excluded from our analysis an additional 108 individuals
that could not be identified with reasonable certainty; most of which were
species of toothed Nomada (73; this group is in need of revision), Osmia males
showing morphological traits of more than 1 common species (18), unknown individuals
from various other genera (9), and damaged specimens (8). We generated
a species-accumulation curve (n = 270, pooled samples from all 30 dunes for all
sampling events in both years) that showed Sorenson distances between subsamples
declining appreciably after just 50 sampling events and confidence intervals collapsing
at 200 sampling events. The species curve had a long tail characteristic of
the presence of singletons in the samples.
Of the 121 species collected and identified, 88 (73%) were represented by fewer
than 10 individuals, and 31 (26%) were represented by only a single individual.
The 5 most-dominant species represented nearly half (48%) of the total number
of specimens and included the sweat bee Lasioglossum floridanum (851 individuals;
noteworthy because this species is primarily southern in its distribution), the
mason bee Osmia pumila (411 individuals), the small carpenter bee Ceratina
calcarata (394 individuals), the sweat bee L. subviridatum (376 individuals),
and the mason bee Osmia sandhouseae (298 individuals). These dominant bees
include ground-nesters (L. floridanum), cavity- and stem-nesters (O. pumila and
C. calcarata) and log-nesters (L. subviridatum) (Cane 1994; Cane et al. 2007;
Gibbs 2011; Medler 1967a, b; Rehan and Richards 2010). The nesting biology of
O. sandhouseae is unknown.
Annual variation
We found 5 species more commonly in 2008 and 4 species in 2009 (Table 1).
Lasioglossum oblongum, a log-nester, was collected only in 2009 (represented by
47 females).
Table 1. Indicators-species analysis (ISA) results for bees exhibiting significant annual variation.
Comparisons are based on the observed indicator values (IV) for species versus values for randomized
groups (1000 randomizations).
Species Max group Observed IV Mean SD P
Agapostemon splendens 2008 15.1 7.1 1.52 0.001
Augochlora pura 2008 29.2 17.8 2.33 0.001
Ceratina strenua 2009 19.9 14 2.04 0.015
Colletes inaequalis 2009 11.2 6.4 1.45 0.007
Habropoda laboriosa 2009 18.5 14.1 1.96 0.038
Lasioglossum bruneri 2008 22.6 12.9 2.12 0.001
Lasioglossum coeruleum 2008 10.1 5.6 1.33 0.006
Lasioglossum oblongum 2009 19.3 7.1 1.46 0.001
Lasioglossum tegulare 2008 22.6 15.9 2.11 0.009
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Seasonal variation
As expected, there were significant seasonal differences in the bee fauna
across the sampling periods: spring, summer, and fall (Table 2). We collected
some species exclusively in only 1 season, while others were collected in 2 or
more seasons but exhibited higher abundance in 1 or more seasons. Appendix 1
lists the exclusive or dominant season (Max group) in which each species was
collected. We report indicator species analysis (ISA) results for all species regardless
of the numbers collected because seasonality is largely phylogenetically
constrained and thus significance is not necessarily dependent on the number of
individuals in the sample. We also report the nesting preferences of each species
(if known or presumed) and the social structure (if known or presumed) based on
available literature in order to look at seasonality data within the context of bee
natural history. The large majority of bee species collected during this study are
ground nesters (69%), followed by those that utilize hollow or pithy stems (12%),
existing cavities (9%), rotten logs (4%), a combination of stems and cavities
(3%), unknown nesting sites (3%), or external or surface nests (less than 1%). In terms of
social structures represented in our samples, 58% of the species we collected are
solitary, 23% are primitive eusocial or semisocial species, 19% are cleptoparasites
or social parasites, and >1% are advanced eusocial (represented by only 1
species, the Western Honey Bee).
There were no family-level seasonal trends evident from our results, although
genus-level trends were apparent, many of which were expected based on prior research
(see Hurd 1979, Michener 2007, references noted in Appendix 2). The ISA
did not indicate significant trends for most species, mainly because so few individuals
were collected for many taxa; significant trends are indicat ed in Appendix 1.
Habropoda laboriosa (Southeastern Blueberry Bee) and all species of Hoplitis
and Nomada were collected only in the spring. Andrena were also collected
exclusively in the spring with the exception of Andrena asteroides, which is a
well-established autumnal species (LaBerge 1967). Similarly, nearly all Osmia
spp. complete their development in the fall and almost always emerge in the
spring; thus, we detected them only in the spring, with the exception of a single
Osmia sandhouseae individual, which was collected in early summer on 7–8 July
2009. All Melissodes were collected in the summer. Other genera were less limited
Table 2. Multiple-response permutation procedure (MRPP) results for seasonal variation. The table
reports the test statistic (T) describing the separation between groups, a measure of effect size (A)
describing within-group agreement, and a P-value representing the likelihood of finding an equal or
smaller delta than the observed based on all possible partitions of the dataset using the Pearson Type
III distribution of deltas.
Groups T A P
Overall -21.360 0.040 less than 0.001
Spring vs. summer -19.197 0.033 less than 0.001
Spring vs. fall -15.090 0.024 less than 0.001
Summer vs. fall -7.262 0.025 less than 0.001
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in terms of flight period; Megachile, Sphecodes, and Lasioglossum all showed
variation in their abundance amongst seasons. Most Lasioglossum exhibited peak
abundance in the summer or fall.
Discussion
Dune habitats likely contain additional species that were not documented in
our study and those that are very rare, and therefore, infrequently encountered, as
well as species that are better represented by collecting methods not employed in
our study (e.g., netting or blue-vane traps) (Cane et al. 2000, Geroff et al. 2014,
Gibbs et al. 2017, Joshi et al. 2015). As other studies have shown (McCravy et
al. 2016, Williams et al. 2001), collecting in different years and seasons helps to
maximize the number of species recorded, especially in terms of documenting the
presence of rare or infrequently collected species. Annual variation may depend on
temperature, rainfall, increased or decreased predation or parasitism, or changes in
resource availability (Cane and Payne 1993, Richards and Packer 1995, Roulston
and Goodell 2011, Tuell and Isaacs 2010). Our records were also determined, in
part, by chance because our traps were deployed only on certain dates and only for
a 48-h period. We know that some species exhibit significant annual variation; thus,
increased collection effort across multiple years and on multiple dates may serve to
increase the number of species documented, especially rare species.
Seasonal variation is largely phylogenetically constrained (Michener 2007).
Pollen specialization may further constrain flight seasons if suitable hosts flower for
a limited period (Fowler 2016). For example, bees exhibiting a preference for Vaccinium
spp. (blueberries), including Andrena bradleyi, Colletes validus, Habropoda
laboriosa, and Osmia virga, were collected only in the spring when these hosts were
in flower. Cleptoparasitic bees necessarily exhibit similar flight times as their hosts,
as was the case with Nomada and their Andrena hosts (Miliczky and Osgood 1995,
Osgood 1989) and with Stelis lateralis and their Hoplitis hosts (Graenicher 1905,
Medler 1961, Michener 1955); we trapped these cleptoparasites only in the spring,
when their hosts were active.
We collected some species of bees throughout the seasonal gradient, although
most peaked in abundance in 1 or more seasons. This pattern is typical of social (e.g.,
Bombus and Lasioglossum [Dialictus]) and occasionally solitary taxa (e.g., Augochlora
pura). These trends were most evident in species that were abundant in our
samples, including many species of Lasioglossum. Some species were collected in all
3 seasons but peaked in abundance in the summer (e.g., L. floridanum, L. fuscipenne,
L. oblongum, L. pectorale, L. subviridatum) or the fall (e.g., L. bruneri, L. pilosum,
L. tegulare, and L. vierecki). Lasioglossum arantium was completely absent from
spring collections, and L. coeruleum was completely absent from fall collections. The
results appear to be independent of social structure or nesting substrate.
Other species that were relatively abundant in our study and collected
throughout the sampling period in all 3 seasons included all species of Ceratina
(C. calcarata and C. strenua reached peak abundance in the summer, C. dupla in
the fall), Agapostemon splendens (summer), Augochlora pura (fall), Augochlorella
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aurata (summer), and Augochloropsis metallica (summer). The results appear to
be independent of social structure. Ceratina may be primarily solitary, although
some females (mother–daughter and daughter–daughter) have a division of labor
and share a nest or nests, and are considered subsocial (Michener 2007, Rehan and
Richards 2010). Ceratina calcarata can also have dwarf, non-reproductive eldest
daughters that forage for their mothers as in eusocial species (Lawson et al. 2016).
Agapostemon splendens and A. pura are solitary. Augochloropsis metallica are most
likely semisocial or eusocial; there is little published information on the nesting biology
of this species (but see Gibbs 2017). The flight period for many species may
be broader than what we documented in our study because our results may have
been limited by rarity of the species, collection methods, or both.
Habitat specialization
Although many bees are pollen specialists and may be restricted in their distribution
based on their host-plant distribution (e.g., Davis and LaBerge 1975,
Eickwort et al. 1986), relatively few bees are known to be narrowly restricted to
a specialized habitat type. Xeric, sandy soil habitats like inland dune and ridge
woodlands may be one of the few examples of specialized habitats that are a
limiting factor for specialist bees (Droege et al. 2009). The reason for the specialization
of these apparently habitat-restricted species is unknown, but it may be
related to both the presence and abundance of specialist plants (J. Selfridge and C.
Frye, unpubl. data; Sorrie 2011) and on the sandy soil substrates (Cane 1991) that
may be required for nesting. The specific characteristics used to select nest sites
are unclear for most bee species.
Several of the bee species we collected are known sand specialists. However,
there are differing degrees of habitat specialization, and discerning the level of
specialization for rare or uncommonly collected species is difficult. Most of the
sand-loving bee species we collected are fairly widespread and occur in a variety
of habitat types as long as sandy soil is a component. These include Agapostemon
splendens and its assumed nest parasite Nomada rubicunda (S. Droege, pers.
comm.), as well as the solitary, ground-nesting Lasioglossum vierecki. Other
species of Lasioglossum assumed to be ground nesters, including L. floridanum,
L. raleighense, and L. sopinci, are also found in areas with loose sandy soil, but
may differ in their level of specialization (e.g., Knerer 1969, Michener 1974, Packer
1993, Sakagami and Michener 1962). Although these species are not as widespread
as A. splendens and L. vierecki, they exhibit some level of habitat flexibility and
again, may not require specialized habitats. Lasioglossum raleighense may be the
most specialized of the 3 taxa, with a majority of specimens collected from GA to
NC, including the Sandhills Region. The Maryland sand dune records are significant
because they represent the northernmost record for the species and, like the
Sandhills Region, they are considered unique, specialized natural communities. The
Sandhills are xeric habitats that are adapted to fire and which support many specialist
plants and animals (Sorrie 2011). Although L. raleighense is not strictly limited
to inland dune and ridge woodland habitats, it certainly appears to exhibit some
level of habitat specialization. Lasioglossum sopinci is another species that occurs
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both in dune habitats and in the Sandhills, but in Maryland has also been recorded
from sandy, non-specialized habitats on the coastal plain (S. Droege, unpubl. data).
Lasioglossum floridanum was the most frequently collected species in our study,
and interestingly, has not been collected anywhere else in the state to date. The reasons
for the absence of previous records are unknown but suggest that there is an
element to this rare community that provides a resource for the species that other
sandy areas throughout the state do not.
The only species we collected that can be linked exclusively to inland dune and
ridge woodlands in the mid-Atlantic region is Lasioglossum arantium, which has
only been collected in dune habitats in Maryland and New Jersey (Gibbs 2011) and
illustrates the importance of these rare natural communities to this apparent habitatspecialist.
Why sand-loving species differ so dramatically in their degree of habitat
specialization is a mystery that we can only piece together with continued study of
species’ life histories. Inventories of bees and other insects in specialized habitats
remain a critical first step in the process of increasing our understanding of these
organisms and their habitats.
Acknowledgments
We are especially thankful to Sam Droege for reviewing early drafts of the manuscript,
sharing data on Maryland bee distribution, and providing assistance with species identification.
We thank the land managers of Chesapeake Forest, Pocomoke State Forest, Shad
Landing State Park, and The Nature Conservancy for allowing us to conduct surveys on
those properties, and Paula Becker, Dana Limpert, Amanda Accamando, Andy Kough,
Sarah Majerowicz, and numerous DNR volunteers for assisting with data collection and
specimen processing. This project was funded through a State Wildlife Grant administered
by the USFWS.
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Appendix 1. List of all bee species collected in 2008 and 2009 across 30 inland dune and ridge woodland sites in Worchester County, MD. Nests are
categorized as ground burrows (G), existing cavities (C), hollow or pithy stems (S), external/surface nests (EX) and rotten logs (W). Cleptoparasites are
indicated using parentheses under the “nests” heading. Social behaviors are categorized as advanced eusocial (A), primitively eusocial or semisocial (E),
solitary to communal (S), and cleptoparasites and social parasites (P). In the highly variable genus Lasioglossum, species with published nesting studies
are marked with an asterisk (*) under the “nests” heading; all others are predicted based on phylogenetic data (Gibbs et al. 2012). Peak abundance evident
from our data (Max group) is described as spring (SPR), summer (SUM), fall (FAL) or (N/A) if peak abundance is the same in all 3 seasons; an asterisk (*)
indicates a significant difference between seasons. For species in which we detected a significant difference, seasons with the same letter in the same row
are not significantly different. Numbers in parenthesis denote actual number of bees collected in a given season throughout the sampling period. A designation
of (ns) denotes that the difference between seasons was not significant. Comparison of abundance across seasons takes into account the maximum
number of sampling events for each season, determined by the number of dunes surveyed (30) multiplied by the number of sampling events in each season
per year (150 spring, 60 summer, 60 fall total).
Observed
Social Max indicator Total # Total #
Species Nests Behavior group value SPR SUM FAL males females
Agapostemon (Agapostemon) splendens (Lepeletier) G S SUM 9.3 ns (15) ns (27) ns (25) 2 65
Agapostemon (Agapostemon) virescens (Fabricius) G S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Andrena (Callandrena) asteroides Mitchell G S FAL 1.7 ns (0) ns (0) ns (1) 1 0
Andrena (Archiandrena) banksi Malloch G S SPR 2.0 ns (6) ns (0) ns (0) 0 6
Andrena (Conandrena) bradleyi Viereck G S SPR 1.3 ns (2) ns (0) ns (0) 2 0
Andrena (Melandrena) carlini Cockerell G S SPR 4.7 ns (9) ns (0) ns (0) 2 7
Andrena (Andrena) cornelli Viereck G S SPR 1.3 ns (2) ns (0) ns (0) 0 2
Andrena (Holandrena) cressonii Robertson G S SPR 2.0 ns (3) ns (0) ns (0) 2 1
Andrena (Scrapteropsis) fenningeri Viereck G S SPR 4.0 ns (7) ns (0) ns (0) 2 5
Andrena (Trachandrena) forbesii Robertson G S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Andrena (Melandrena) hilaris Smith G S SPR 1.3 ns (2) ns (0) ns (0) 2 0
Andrena (Scrapteropsis) imitatrix Cresson G S SPR 2.7 ns (5) ns (0) ns (0) 0 5
Andrena (Larandrena) miserabilis Cresson G S SPR* 10.7 (28)A (0)B (0)B 1 27
Andrena (Andrena) tridens Robertson G S SPR 2.7 ns (4) ns (0) ns (0) 0 4
Andrena (Melandrena) vicina Smith G S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Andrena (Lomelissa) violae Robertson G S SPR 4.7 ns (9) ns (0) ns (0) 5 4
Anthidiellum (Loyolanthidium) notatum (Latreille) EX S FAL 3.3 ns (0) ns (0) ns (2) 0 2
Apis (Apis) mellifera L.1 C A FAL 4.8 ns (3) ns (5) ns (7) 0 15
Augochlora (Augochlora) pura (Say) W S FAL* 31.8 (12)A (89)B (99)B 21 179
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Observed
Social Max indicator Total # Total #
Species Nests Behavior group value SPR SUM FAL males females
Augochlorella aurata (Smith) G E SUM* 22.9 (23)A (83)B (76)B 2 180
Augochloropsis (Paraugochloropsis) metallica fulgida (Smith) G E SUM* 7.6 (3)A (7)B (1)AB 0 11
Augochloropsis (Paraugochloropsis) metallica metallica (Fabricius) G E SUM* 6.1 (1)A (4)B (0)AB 1 4
Bombus (Pyrobombus) bimaculatus Cresson C E SPR/SUM 2.4 ns (1) ns (1) ns (0) 0 2
Bombus (Thoracobombus) fervidus (Fabricius) C E SPR 0.7 ns (1) ns (0) ns (0) 0 1
Bombus (Cullumanobombus) griseocollis (DeGeer) C E SPR 0.7 ns (1) ns (0) ns (0) 0 1
Bombus (Pyrobombus) impatiens Cresson C E FAL* 7.7 (6)A (5)AB (10)B 4 11
Bombus (Thoracobombus) pensylvanicus (DeGeer) C E FAL 1.7 ns (0) ns (0) ns (2) 0 2
Calliopsis (Calliopsis) andreniformis Smith C S FAL 3.3 ns (0) ns (0) ns (3) 0 3
Ceratina (Zadontomerus) calcarata Robertson S S SUM* 34.7 (71)A (201)B (122)C 194 200
Ceratina (Zadontomerus) dupla Say S S SUM 6.4 ns (2) ns (14) ns (11) 2 25
Ceratina (Zadontomerus) strenua Smith S S SUM* 29.1 (29)A (132)B (35)A 71 125
Coelioxys (Xerocoelioxys) immaculatus Cockerell (G) P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Coelioxys (Boreocoelioxys) porterae Cockerell C P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Colletes americanus Cresson G S FAL 1.7 ns (0) ns (0) ns (2) 2 0
Colletes inaequalis Say G S SPR* 16.7 (48)A (0)B (0)B 3 45
Colletes thoracicus Smith G S SPR 4.0 ns (12) ns (0) ns (0) 1 11
Colletes validus Cresson G S SPR 4.7 ns (9) ns (0) ns (0) 5 4
Epeolus pusillus Cresson (G) P FAL 1.7 ns (0) ns (0) ns (1) 1 0
Eucera hamata (Bradley) G S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Habropoda laboriosa (Fabricius) G S SPR* 42.0 (270)A (0)B (0)B 178 92
Halictus (Odontalictus) ligatus Say/H. (O.) poeyi Lepeletier2 G E FAL 2.0 ns (1) ns (2) ns (5) 4 4
Halictus (Protohalictus) rubicundus (Christ) G E SPR 1.3 ns (2) ns (0) ns (0) 0 2
Heriades (Neotrypetes) variolosa (Cresson) S S FAL 1.7 ns (0) ns (0) ns (1) 0 1
Hoplitis (Alcidamea) pilosifrons (Cresson) S S SPR 2.7 ns (4) ns (0) ns (0) 4 0
Hoplitis (Alcidamea) producta (Cresson) S S SPR 3.3 ns (5) ns (0) ns (0) 1 4
Hoplitis (Alcidamea) spoliata (Provancher) S S SPR 2.0 ns (3) ns (0) ns (0) 1 2
Hylaeus (Prosopis) affinis (Smith) S S FAL 1.7 ns (0) ns (0) ns (1) 0 1
Hylaeus (Prosopis) modestus Say S S N/A 0.7 ns (1) ns (1) ns (1) 2 1
Hylaeus (Metziella) sparsus (Cresson) S S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Lasioglossum (Dialictus) arantium Gibbs G E SUM* 21.6 (0)A (154)B (60)C 7 207
Lasioglossum (Hemihalictus) birkmanni (Crawford) G S FAL 5.6 ns (2) ns (0) ns (3) 0 5
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Observed
Social Max indicator Total # Total #
Species Nests Behavior group value SPR SUM FAL males females
Lasioglossum (Dialictus) bruneri (Crawford) G E FAL* 20.1 (4)A (61)B (73)B 4 134
Lasioglossum (Dialictus) coeruleum (Robertson) W E* SUM* 9.8 (10)A (18)AB (0)B 2 26
Lasioglossum (Dialictus) coreopsis (Robertson) G E SPR 4.9 ns (8) ns (3) ns (6) 0 17
Lasioglossum (Dialictus) cressonii (Robertson) W E FAL 4.4 ns (1) ns (0) ns (3) 0 4
Lasioglossum (Dialictus) floridanum (Robertson) G E SUM* 31.6 (173)A (488)B (190)A 14 837
Lasioglossum (Hemihalictus) foxii (Robertson) G S SUM 3.3 ns (0) ns (2) ns (0) 2 0
Lasioglossum (Lasioglossum) fuscipenne (Smith) G S SUM 5.3 ns (5) ns (8) ns (5) 3 15
Lasioglossum (Dialictus) hitchensi Gibbs 2012 G E SPR 1.9 ns (4) ns (2) ns (0) 0 6
Lasioglossum (Dialictus) illinoense (Robertson) G E FAL 1.7 ns (0) ns (0) ns (1) 0 1
Lasioglossum (Dialictus) imitatum (Smith) G E* FAL 1.1 ns (0) ns (1) ns (2) 0 3
Lasioglossum (Hemihalictus) lustrans (Cockerell) G S* SUM 3.3 ns (0) ns (2) ns (0) 0 2
Lasioglossum (Hemihalictus) nelumbonis (Robertson) G S SPR/SUM 1.9 ns (1) ns (1) ns (0) 0 2
Lasioglossum (Dialictus) oblongum (Lovell) W E SUM* 9.9 (15)AB (28)A (4)B 0 47
Lasioglossum (Hemihalictus) pectorale (Smith) G S SUM* 15.9 (3)A (16)B (1)A 2 18
Lasioglossum (Dialictus) pilosum (Smith) G E FAL* 10.4 (4)A (13)AB (23)B 3 38
Lasioglossum (Dialictus) platyparium (Robertson) G P SPR/FAL 1.2 ns (1) ns (0) ns (1) 0 2
Lasioglossum (Dialictus) raleighense (Crawford) G E SPR/FAL 0.7 ns (1) ns (0) ns (1) 0 2
Lasioglossum (Dialictus) smilacinae (Robertson) G E SUM 2.8 ns (0) ns (2) ns (1) 0 3
Lasioglossum (Hemihalictus) sopinci (Crawford) G S SUM 5.6 ns (6) ns (9) ns (2) 0 17
Lasioglossum (Dialictus) subviridatum (Cockerell) W E SUM* 53.5 (94)A (275)B (7)C 0 376
Lasioglossum (Dialictus) tegulare (Robertson) G E FAL* 21.9 (33)A (35)AB (91)B 3 156
Lasioglossum (Dialictus) versatum (Robertson) G E* FAL* 20.1 (4)A (0)AB (5)B 1 8
Lasioglossum (Dialictus) vierecki (Crawford) G S* FAL 12.9 ns (21) ns (78) ns (84) 5 178
Lasioglossum (Dialictus) weemsi (Mitchell) G E SUM 1.7 ns (0) ns (3) ns (0) 0 3
Lasioglossum (Dialictus) zephyrum (Smith) G E* SUM/FAL 0.8 ns (0) ns (1) ns (1) 0 2
Megachile (Litomegachile) brevis Say S/C S FAL 1.7 ns (0) ns (0) ns (1) 0 1
Megachile (Chelostomoides) exilis Cresson C S FAL 1.7 ns (0) ns (0) ns (1) 0 1
Megachile (Xanthosarus) gemula Cresson S/C S SPR 1.3 ns (2) ns (0) ns (0) 1 1
Megachile (Litomegachile) mendica Cresson S/C S SUM 4.4 ns (1) ns (3) ns (0) 4 0
Megachile (Xanthosarus) mucida Cresson G S SPR 0.7 ns (1) ns (0) ns (0) 1 0
Megachile (Litomegachile) texana Cresson G S SUM 3.3 ns (0) ns (1) ns (0) 0 1
Melissodes (Melissodes) bimaculatus (Lepeletier) G S SUM 3.3 ns (0) ns (2) ns (0) 0 2
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Observed
Social Max indicator Total # Total #
Species Nests Behavior group value SPR SUM FAL males females
Melissodes (Melissodes) comptoides Robertson G S SUM* 8.3 (0)A (5)B (0)A 2 3
Melissodes (Eumelissodes) denticulatus Smith G S SUM 3.3 ns(0) ns (2) ns (0) 0 2
Melissodes (Eumelissodes) druriellus (Kirby) G S SUM 5.0 ns (0) ns (3) ns (0) 1 2
Melissodes (Eumelissodes) trinodis Robertson G S SUM* 11.7 (0)A (10)B (0)A 0 10
Nomada armatella Cockerell (G) P SPR 4.7 ns 10) ns (0) ns (0) 8 2
Nomada articulata Smith (G) P SPR 2.7 ns (7) ns (0) ns (0) 6 1
Nomada composita Mitchell (G) P SPR 1.3 ns (2) ns (0) ns (0) 0 2
Nomada cressonii Robertson (G) P SPR 4.0 ns (6) ns (0) ns (0) 6 0
Nomada denticulata Robertson (G) P SPR 1.3 ns (2) ns (0) ns (0) 2 0
Nomada luteoloides Robertson (G) P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Nomada luteola Olivier (G) P SPR 3.3 ns (4) ns (0) ns (0) 4 0
Nomada maculata Cresson (G) P SPR 3.3 ns (8) ns (0) ns (0) 6 2
Nomada parva Robertson (G) P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Nomada pygmaea Cresson (G) P SPR* 8.0 (9)A (0)B (0)B 6 3
Nomada rubicunda Olivier (G) P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Nomada sulphurata Smith (G) P SPR 0.7 ns (1) ns (0) ns (0) 1 0
Nomia (Acunomia) maneei Cockerell G S SUM* 6.7 (0)A (6)B (0)AB 1 5
Osmia (Melanosmia) atriventris Cresson S S SPR* 41.3 (181)A (0)B (0)B 130 51
Osmia (Melanosmia) collinsiae Robertson ? S SPR 1.3 ns (3) ns (0) ns (0) 3 0
Osmia (Helicosmia) georgica Cresson S S SPR 1.3 ns (2) ns (0) ns (0) 1 1
Osmia (Melanosmia) inspergens Lovell and Cockerell S S SPR* 16.7 (69)A (0)B (0)B 67 2
Osmia (Osmia) lignaria lignaria Say S S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Osmia (Melanosmia) pumila Cresson S S SPR* 45.3 (411)A (0)B (0)B 373 38
Osmia (Melanosmia) sandhouseae Mitchell ? S SPR* 39.0 (297)A (1)B (0)B 273 25
Osmia (Osmia) taurus Smith S S SPR 3.3 ns (5) ns (0) ns (0) 4 1
Osmia (Melanosmia) virga Sandhouse ? S SPR* 24.7 (62)A (0)B (0)B 43 19
Panurginus atramontensis Crawford G S SPR 0.7 ns (1) ns (0) ns (0) 0 1
Peponapis (Peponapis) pruinosa (Say) G S SUM 3.3 ns (0) ns (3) ns (0) 2 1
Perdita (Alloperdita) bradleyi Viereck G S SPR 1.3 ns (3) ns (0) ns (0) 0 3
Perdita (Perdita) octomaculata (Say) G S FAL 5.0 ns (0) ns (0) ns (3) 1 2
Sphecodes aroniae Mitchell (G) P SPR 0.7 ns (1) ns (0) ns (0) 0 1
Sphecodes banksii Lovell (G) P FAL 2.8 ns (1) ns (0) ns (2) 2 1
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Observed
Social Max indicator Total # Total #
Species Nests Behavior group value SPR SUM FAL males females
Sphecodes confertus Say (G) P SPR 1.3 ns (2) ns (0) ns (0) 0 2
Sphecodes coronus Mitchell (G) P SPR 0.7 ns (1) ns (0) ns (0) 0 1
Sphecodes cressonii (Robertson) (G) P SPR 2.7 ns (4) ns (0) ns (0) 0 4
Sphecodes mandibularis Cresson (G) P SPR 0.7 ns (1) ns (0) ns (0) 0 1
Sphecodes fattigi Mitchell (G) P SUM 1.7 ns (0) ns (1) ns (0) 0 1
Stelis (Stelis) lateralis Cresson (S) P SPR 0.7 ns (1) ns (0) ns (0) 0 1
Svastra (Epimelissodes) obliqua (Say, 1837) G S SUM 1.7 ns (0) ns (1) ns (0) 0 1
1Introduced
2The two species cannot currently be separated morphologically (Packer et al. 2016).
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Appendix 2. Literature cited summary for each genus listed in Appendix 1.
Genus Literature Cited
Agapostemon Abrams and Eickwort 1980, Eickwort 1981
Andrena Miliczky 1988, Miliczky and Osgood 1995, Norden and Scarborough 1979,
Osgood 1989, Schrader and LaBerge 1978
Anthidiellum Baker et al. 1985
Apis Michener 1974
Augochlora Stockhammer 1966
Augochlorella Mueller 1996, Ordway 1966, Packer 1990
Augochloropsis Eickwort and Sakagami 1979, Gibbs 2017
Bombus Michener 1974
Calliopsis Shinn 1967
Ceratina Kislow 1976, Lawson et al. 2016, Rehan and Richards 2010, Vickruck et al.
2011
Coelioxys Baker 1975
Colletes Batra 1980, Rajotte 1979
Epeolus Rozen and Favreau 1968
Eucera Miliczky 1985
Habropoda Cane 1994
Halictus Dunn et al. 1998, Richards and Packer 1995, Richards et al. 2010, Soucy 2002
Heriades Fischer 1955
Hoplitis Fye 1965; Medler 1961, 1967a; Michener 1955
Hylaeus Krombein 1967, Rau 1930
Lasioglossum Batra 1964; Breed 1975; Daly 1961; Gibbs 2011; Gibbs et al. 2012a, 2012b,
2013; Michener 1947; Michener and Wille 1961; Mitchell 1960; Sakagami
and Michener 1962; Stockhammer 1967
Megachile Krombein 1953, Medler 1965, Michener 1953, Sheffield et al. 201 1
Melissodes Ashmead 1894, Cameron et al. 1996, Clement 1973, Graenicher 1905
Nomada Abrams and Eickwort 1981, Goldstein and Ascher 2016, Miliczky and Osgood
1995, Osgood 1989
Nomia Cross and Bohart 1960
Osmia Fye 1965, Hartman et al. 1944, Krombein 1967, Hogendorn and Leys 19987,
Medler 1967b, Medler and Lussenhop 1968
Panurginus Rozen 1967
Peponapis Hurd et al. 1974
Perdita Eickwort 1977
Sphecodes Michener 1978
Stelis Graenicher 1905, Medler 1961, Michener 1955
Svastra Rozen 1964