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2019 THE GREAT LAKES ENTOMOLOGIST 1
The status of wild bee species and
populations has been the subject of a great
deal of attention by the scientic community
in recent years, with rapidly contracting
distributions for bees such as Bombus afnis
Cresson documented in the United States
and Canada (Colla and Packer 2008; Cam-
eron et al. 2011). Understanding population
declines in wild bees outside of the genus
Bombus has been more difcult, in part be-
cause many solitary bee species are rare in
collections, either because they have small
geographic ranges or because they are phe-
nologically limited to a narrow ight period
and require specialized collection effort in
order to detect (Harrison et al. 2017). Indeed,
in a study of the population trends of wild
bees in the northeastern United States, 87
of the 438 species (19.9%) were represented
by only 10 to 30 specimens over a 140-year
period, making assessment of their historic
and contemporary statuses challenging
(Bartomeus et al. 2013).
One bee species that has been very
infrequently recorded in North America is
Epeoloides pilosulus (Cresson). Epeoloides
pilosulus is a brood parasite of Macropis bees
(Shefeld et al. 2004; Wagner and Ascher
2008), which are themselves specialists on
Lysimachia species (Fig. 1A, Primulaceae),
collecting pollen and oral oils exclusively
from this genus (Michez and Patiny 2005).
Floral oils are mixed with pollen provisions
and used to waterproof the linings of the cell
wall (Cane et al. 1983), allowing Macropis to
nest in the damp soils favored by Lysimachia
species (Fig. 1B). Epeoloides pilosulus is thus
twice restricted; rst by the limited suite of
bee species that it parasitizes and second
by the narrow ecological niche occupied by
its hosts.
Epeoloides pilosulus is consequently
very rarely collected. Most specimens in
Michigan were collected in the early part of
the 20th century with the last record made
in 1944. This ts into the overall trend for
E. pilosulus, as the species was not recorded
in North America between 1960 and 2002
(but see Shefeld and Heron 2018), until
it was rediscovered in Nova Scotia based
on two male specimens collected in a pan
trap (Shefeld et al. 2004). There have only
been four additional contemporary records
of E. pilosulus in North America since then,
in Connecticut (2006, Wagner and Ascher
2008), Alberta (2010, Shefeld and Heron
2018), New York (2014, http://bugguide.
net/node/view/954741), and Maine (2016,
Dibble et al. 2017). In Michigan, E. pilosulus
is known from four counties in the central
and southern Lower Peninsula (Berrien,
Midland, Van Buren, and Wayne).
Epeoloides pilosulus
(Cresson) (Hymenoptera: Apidae)
Rediscovered in Michigan, with Notes on the
Distribution and Status of its
Macropis
Hosts
T.J. Wood1,2*, M.F. Killewald1,3, K.K. Graham1, J. Gibbs3 and R. Isaacs1
1 Department of Entomology, Michigan State University, East Lansing, MI, 48824, USA
2 Current afliation: Laboratory of Zoology, University of Mons, 7000, Mons, Belgium
3 Department of Entomology, University of Manitoba, 12 Dafoe Rd.,
Winnipeg, MB, R3T 2N2, Canada
*Corresponding author: (e-mail: thomasjames.wood@umons.ac.be)
Abstract
Epeoloides pilosulus (Cresson 1878) (Hymenoptera: Apidae) is one of the rarest bees
in North America with only a handful of records since 1960. Epeoloides pilosulus is a brood
parasite of Macropis bees, which until recently had not been collected in Michigan since
1944. Bee surveys in Midland County, Michigan have led to the rediscovery of E. pilosulus
in this state – the rst record in 74 years. Michigan becomes the fourth state where E. pi-
losulus has been rediscovered after Connecticut in 2006, New York in 2014 and Maine in
2016, and the sixth region in North America after Nova Scotia in 2002 and Alberta in 2010.
State-wide bee surveys have also shown that the principal host, Macropis nuda (Provancher
1882), remains widespread in Michigan, and that Macropis patellata Patton 1880 is newly
recorded for the state.
Key words: parasitic bee, oil bee, oligolege, bee surveys, Lysimachia
1
Wood et al.: Epeoloides pilosulus rediscovered in Michigan
Published by ValpoScholar, 2019
2 THE GREAT LAKES ENTOMOLOGIST Vol. 52, Nos. 1–2
Only one species of Macropis has been
recorded from Michigan – Macropis nuda
(Provancher). Macropis nuda has a large
range and is known from Canada, from
British Columbia to Nova Scotia, and the
United States, from Montana and Colorado
to the New England states (Mitchell 1960;
Michez and Patiny 2005; Shefeld and Heron
2018). However, like E. pilosulus, M. nuda
appeared to have disappeared from Mich-
igan, being last recorded in 1959 and not
rediscovered until 2017 in Hillsdale County
(Gibbs et al. 2017).
The aim of this paper is to report on the
ndings of recent bee faunal surveys across
Michigan that include the rediscovery of E.
pilosulus and have expanded our under-
standing of the status of Macropis species
in the state.
Figure 1. A. Fringed Loosestrife (Lysimachia ciliata L.) ower. B. Lysimachia ciliata in ower in damp
prairie fen habitat at Ives Road Fen, Lenawee County. C. European Dotted Loosestrife (Lysimachia
punctata L.) outside an abandoned house in Felch, Dickinson County. D-F. Macropis nuda (Provancher)
individuals at Algonac State Park, St. Clair County. D. Macropis nuda male showing distinctive yellow
facial maculations. E-F. Macropis nuda female collecting pollen from L. ciliata.
2
The Great Lakes Entomologist, Vol. 52, No. 1 [2019], Art. 4
https://scholar.valpo.edu/tgle/vol52/iss1/4
2019 THE GREAT LAKES ENTOMOLOGIST 3
Methods
As part of our Michigan bee survey
we visited the Averill Preserve (43.6618,
–84.3500; managed by Little Forks Conser-
vancy) in Midland County every other week
from mid-June to mid-September in 2017
and 2018. During each visit, we spent one
cumulative hour sampling bees using aerial
nets, and we recorded which plants bees were
caught on. Surveys focused on open habitats,
with surveyors searching all owering plants
within these areas. Similar sampling meth-
ods were used at seven other sites in Mid-
land, Ingham, Shiawassee, Kalamazoo, and
Livingston counties. All bees were pinned
and labelled at Michigan State University,
and then identied to species at the Uni-
versity of Manitoba (JG and Joel Gardner).
During collection at the Averill Preserve, a
specimen of E. pilosulus was captured (see
Results). This material is deposited in the
Albert J. Cook Arthropod Research Collec-
tion at Michigan State University.
The records of Macropis species for
Michigan stated in this paper are based on
the collections of the lead author as part of
his efforts to survey wild bees in every county
in Michigan. His material is deposited at
the J.B. Wallis / R. E. Roughley Museum
of Entomology, University of Manitoba.
Historic records of E. pilosulus and M. nuda
collections were taken from the most recent
summary of the Michigan bee fauna (Gibbs
et al. 2017).
Results
Epeoloides pilosulus (Cresson 1878)
Current records: Midland Co.: Mid-
land, Averill Preserve, 43.6618, –84.3500, M.
Killewald, 28 June 2018, 1♀, (BH_010955),
Lysimachia nummularia L.
The specimen was collected as it was
patrolling a small patch of L. nummularia
owers along a walkway.
Macropis (Macropis) nuda
(Provancher 1882)
Current records: Alcona Co.:
Black River, Black River Road x La Vigne
Road, 44.815, –83.324, T.J. Wood, 15 July
2018, 1♀, Apocynum androsaemifolium L.;
Alpena Co.: Mackinaw State Forest, Long
Rapids Rd x Truax Creek, 45.116, –83.823,
T.J. Wood, 15 July 2018, 1♂, 1♀, Lysima-
chia ciliata L.; Dickinson Co.: Foster City,
Felch, 45.996, –87.825, T.J. Wood, 30 June
2018, 2♂, Lysimachia punctata L., Fig. 2C;
Hillsdale Co.: Pittsford State Game Area,
41.866, –84.522, T.J. Wood, 8 July 2017, 1♂,
Apocynum cannabinum L.; St. Clair Co.:
Algonac State Park, 42.650, –82.531, T.J.
Wood, 14 July 2018, 1♂, 1♀, L. ciliata, Fig.
1D–F; Tuscola Co.: Dayton, S Plain Road
x James Road, 43.462, –83.268, T.J. Wood,
25 June 2018, 1♂, 1♀, L. ciliata.
Macropis nuda was not known from
Alcona, Hillsdale, St. Clair and Tuscola coun-
ties prior to its discovery there in 2017–2018.
The historic and contemporary distributions
are shown in Fig. 2A.
Macropis (Macropis) patellata
Patton 1880
Current records: Lenawee Co.: Ives
Road Fen Preserve, 41.967, –83.945, T.J.
Wood, 8 July 2018, 3♂, L. ciliata.
Males and females were abundant at
an area of restored prairie fen in south-east
Michigan with regenerating wetland vege-
tation (Fig. 2B). No females were collected
as TJW believed at the time that these bees
were M. nuda, and consequently did not col-
lect any females in order to avoid depleting
the population. It was not until they were
inspected under the microscope that their
true identity was determined.
Discussion
The rediscovery of E. pilosulus in
Midland County suggests that the species
has been present in Michigan continuously
since it was rst discovered over a century
ago. Although M. nuda was not also found
at the same site, the host and the parasite
are often detected using different sampling
techniques, with the parasite detected using
bowl traps and the host using aerial netting
in both Nova Scotia and Connecticut (Shef-
eld et al. 2004; Wagner and Ascher 2008).
Contemporary records of M. nuda suggest
that this species remains most common in
the Saginaw Bay region of eastern Michigan
(Fig. 2A).
All pollen foraging M. nuda females
along with several males were collected
from Fringed Loosestrife (L. ciliata). This
is in common with other studies in eastern
North America (Cane et al. 1983). Males
were also collected patrolling around the
introduced European Dotted Loosestrife
(L. punctata) in Dickinson County (Fig.
1C), suggesting that it may also be visited
by females of this species. In Europe, L.
punctata is visited for pollen and oil by the
native M. fulvipes (Fabricius) (Michez and
Patiny 2005). In contrast, the European
Creeping Jenny (L. nummularia) is not vis-
ited by Macropis species (Cane et al. 1983)
as it does not produce oil. This may explain
why E. pilosulus was found at the Averill
Preserve site but M. nuda was not. Surveys
3
Wood et al.: Epeoloides pilosulus rediscovered in Michigan
Published by ValpoScholar, 2019
4 THE GREAT LAKES ENTOMOLOGIST Vol. 52, Nos. 1–2
were restricted to dry, open areas, where no
other Lysimachia species were present, al-
though they may have been present in damp
riverine areas nearby. Individual females
of M. nuda collected on Indian Hemp (A.
cannabinum) and Spreading Dogbane (A.
androsaemifolium) showed no evidence of
pollen collection and were visited solely for
nectar. Though restricted to Lysimachia for
pollen and oral oils, Macropis species have
been recorded visiting many plant families
for nectar (Pekkarinen et al. 2003; Michez
and Patiny 2005), including Apocynum by
M. nuda (Cane et al. 1983).
Macropis patellata is known to occur in
the eastern United States from North Caro-
lina north to Vermont and west to Iowa and
Nebraska (Mitchell 1960; Michez and Patiny
2005; Ascher and Pickering 2018). Surpris-
ingly, the species has never been recorded
from Indiana or Ohio, so this record from
south-eastern Michigan lls a distributional
gap and suggests that, in addition to these
two states, the species may also be present
in southern Ontario. Most of the data used
to determine the distribution of this bee are
old, and the species was highlighted as po-
tentially being of conservation concern due to
a lack of recent records in the northeastern
United States (Bartomeus et al. 2013). The
two other Macropis species known from
North America, M. ciliata Patton 1880 and
M. steironematis Robertson 1891, have a
similar distribution to M. patellata, being
found in some Atlantic states and parts of
the Midwest, but have not been recorded
from Michigan, Indiana or Ohio (Michez and
Patiny 2005; Ascher and Pickering 2018).
Continued targeted searching may reveal the
presence of one or both of these additional
Macropis species in this region.
The rediscovery of M. nuda (Gibbs et
al. 2017) and E. pilosulus, as well as the
discovery of M. patellata, in Michigan high-
lights some potential consequences of low
sampling effort for bees over long periods
of time. These include a high likelihood of
missing bees that have restricted geograph-
ical ranges or that exploit a narrow range
of host plants, as well as the potential to
consider a bee rare when it has instead been
poorly sampled. The level of active sampling
for bees in Michigan has, until very recently,
been low in comparison with the historical
baseline. The year of the most recent pre-
vious record for M. nuda (1959) coincides
with the nal collection period of R.R. Dreis-
bach, a prolic amateur entomologist who
collected bees extensively across the whole
state (Fischer 1965). Macropis nuda and E.
pilosulus were almost certainly continuous-
ly present in Michigan for the last 60–70
years, and the absence of records for these
species likely reects low sampling effort
rather than a genuine population decline.
More regular and extensive sampling is
needed to inform conservation efforts along
with targeted searches to understand the
Figure 2. A. Distribution of Macropis nuda (Provancher) in Michigan at a county-level resolution. The
single new record of Macropis patellata Patton is marked with an asterisk. B. Distribution of Epeoloides
pilosulus (Cresson) in Michigan at a county-level resolution. Pre-1960 records are marked in gray and
2017–2018 records are marked in red. Hatching indicates records of the species in both time periods.
4
The Great Lakes Entomologist, Vol. 52, No. 1 [2019], Art. 4
https://scholar.valpo.edu/tgle/vol52/iss1/4
2019 THE GREAT LAKES ENTOMOLOGIST 5
abundance and distribution of specialized
species that have narrow ecological niches
which make them difcult to detect. General
bee surveys in North America are likely at
an all-time high, but many species of con-
servation concern may best be studied with
more focused efforts.
Acknowledgments
Survey work was funded by USDA
NIFA grant 2017-68004-26323. Our thanks
go to Gary Parsons for access to the insect
collection at MSU, to The Little Forks Con-
servancy for allowing access to the Averill
site, and to The Nature Conservancy for
allowing access to the Ives Road Fen site.
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