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A Checklist of the Bees (Hymenoptera: Apoidea) of St. Louis, Missouri, USA

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Concern over the declines of pollinator populations during the last decade has resulted in calls from governments and international agencies to better monitor these organisms. Recent studies of bee diversity in urban environments suggest that cities may contain significant amounts of bee species, even greater than surrounding agricultural areas, and in some occasions comparable to natural habitats. We conducted a three-year survey of bees in the city of St. Louis, MO. Like many other post-industrial cities in the United States, St. Louis is considered a shrinking city, with many vacant lots and unoccupied structures, mostly in the urban core. We sampled a broad range of habitats throughout the growing seasons of 2013 to 2016, e.g., vacant lots, city parks, community gardens, and urban farms, using aerial netting. This resulted in over 7,700 specimens. Data from other surveys, e.g., BioBlitz, and personal collections was also utilized in developing the species list. These data were supplemented with inspections of entomological collections from institutions in the state and the scientific literature. We identified a total of 198 species of bees from five different families that occur in the city. Only nine of the bee species present in the city are non-natives. The city of St. Louis currently hosts nearly 45% of the bee diversity of the state, likely making it one of most species-rich cities relative to its state's total bee fauna in the country. This represents a great natural resource that must be better understood, and has potential conservation implications.
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A Checklist of the Bees (Hymenoptera: Apoidea) of St. Louis,
Missouri, USA
Author(s): Gerardo R. Camilo, Paige A. Muñiz, Michael S. Arduser, and Edward
M. Spevak
Source: Journal of the Kansas Entomological Society, 90(3):175-188.
Published By: Kansas Entomological Society
https://doi.org/10.2317/0022-8567-90.3.175
URL: http://www.bioone.org/doi/full/10.2317/0022-8567-90.3.175
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JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
90(3), 2017, pp. 175–188
A Checklist of the Bees (Hymenoptera: Apoidea) of
St. Louis, Missouri, USA
GERARDO R. CAMILO,1,*PAIGE A. MU˜
NIZ,1MICHAEL S. ARDUSER,2AND EDWARD
M. SPEVAK3
ABSTRACT: Concern over the declines of pollinator populations during the last decade has resulted
in calls from governments and international agencies to better monitor these organisms. Recent studies
of bee diversity in urban environments suggest that cities may contain significant amounts of bee
species, even greater than surrounding agricultural areas, and in some occasions comparable to natural
habitats. We conducted a three-year survey of bees in the city of St. Louis, MO. Like many other post-
industrial cities in the United States, St. Louis is considered a shrinking city, with many vacant lots and
unoccupied structures, mostly in the urban core. We sampled a broad range of habitats throughout the
growing seasons of 2013 to 2016, e.g., vacant lots, city parks, community gardens, and urban farms,
using aerial netting. This resulted in over 7,700 specimens. Data from other surveys, e.g., BioBlitz,
and personal collections was also utilized in developing the species list. These data were supplemented
with inspections of entomological collections from institutions in the state and the scientific literature.
We identified a total of 198 species of bees from five different families that occur in the city. Only nine
of the bee species present in the city are non-natives. The city of St. Louis currently hosts nearly 45%
of the bee diversity of the state, likely making it one of most species-rich cities relative to its state’s
total bee fauna in the country. This represents a great natural resource that must be better understood,
and has potential conservation implications.
KEY WORDS: Anthophila, urban biodiversity, urban pollinators, shrinking city, novel ecosystem
Bees are arguably the most important pollinators on a global scale (Buchmann and
Nabhan, 1996). Yet, many species are in trouble (Shepherd et al., 2003; Winfree, 2010).
Declines of honeybees and North American native bees, such as bumblebees, have been
reported over the past decade (Grixti et al., 2009; Williams and Osborne, 2009; Winfree
et al., 2009). The status of many native bees is not well understood, and according to some
has already reached a crisis stage (Dixon, 2009). The dire lack of data is perhaps best
described by the National Academy of Sciences (2007) in a recent report on the status of
pollinators: “...the paucity of long-term data and the incomplete knowledge of even basic
taxonomy and ecology make the definitive assessment of status exceedingly difficult.” It is
estimated that there are over 20,000 species worldwide (Michner, 2000), and around 4000
in the continental United States (Wilson and Carrill, 2015). For the state of Missouri, it is
estimated that there are some 452 species of bees (M. Arduser, unpublished data). Yet, little
is known about the distribution of bees for urban environments in general, and for the city
of St. Louis in particular.
As of the year 2000, approximately half of the world’s population lives in urban areas
(Millennium Ecosystem Assessment, 2005). The same report proposed that by the middle
of the century, two out of every three human beings will be living in an urban environment.
At the same time, the pressure on agricultural systems to produce food for all those
people will be greater than ever. Thus, understanding the mechanisms that maintain and
1Department of Biology, Saint Louis University, 3507 Laclede Ave., St. Louis, MO 63103
2325 Atalanta Ave., St. Louis, MO 63119
3Saint Louis Zoo, One Government Drive, St. Louis, MO 63110
*Corresponding author: camilogr@slu.edu
Received 2 December 2016; Accepted 18 November 2017
C
2017 Kansas Entomological Society
176 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
mediate pollinator diversity and abundance in urban environments is critical (Garibaldi
et al., 2014).
The current approach to native bee conservation is habitat enhancement (Shepherd,
2002; Shepherd et al., 2003; Shepherd et al., 2008). Common recommendations include
providing nesting and foraging resources in developed landscapes in hopes that pockets
of suitable habitat will sustain diverse bee communities. Human population decline in the
urban core of St. Louis city, further exacerbated by widespread foreclosures during the
economic recession of 2008–2010, have created novel ecosystems (Morse et al., 2014)
that hint at potential sites for general conservation (Frazier and Bagchi-Sen, 2015), and
insect pollinators specifically (Gardiner et al., 2013; Burr et al., 2016). Many of these
newly open spaces could be considered enhanced habitat from the perspective of native
bees (Baldock et al., 2015; Threlfall et al., 2015; Hall et al., 2016). Abandoned buildings,
or those in disrepair, may provide cavities for females to nest. Brownfields and open lots
provide undisturbed soils for ground-dwelling species, and weedy vegetation that can be
quite attractive, as typical urban-exploiting plant species such as clover and various asters
are highly attractive to bees. Utilization of a small proportion of these lots by the community
to convert them into parks or community gardens further enhances sites (Frazier and Bachi-
Sen, 2015) and potentially provides more resources for native bee conservation (Burr et al.,
2016; Hall et al., 2016).
Indeed, recent work has proposed that cities may represent a refuge of sorts for many
wild bees (Hall et al., 2016). For example, the bulk of the bee diversity in England resides
in urban environments instead of more natural or agrarian habitats (Baldock et al., 2015).
Less striking, yet still surprisingly high levels of bee diversity have been observed in Berlin,
Germany (Saure et al., 1998), Melbourne, Australia (Threlfall et al., 2015), Guanacaste
Province, Costa Rica (Frankie et al., 2013), Vancouver, Canada (Tommasi et al., 2004),
Chicago, IL (Tonietto et al., 2011; Lowenstein et al., 2014), and New York City, NY
(Matteson et al., 2008; Matteson and Langellotto, 2009).
The objective of our study was to assemble a comprehensive checklist of bee species
for St. Louis city that could serve as a comparison to other urban environments, and as a
reference for further studies in the St. Louis metropolitan area. St. Louis is representative of
the post-industrial, rust-belt, Midwestern city in North America (Gordon, 2008). There has
been tremendous urban sprawl during the last twenty years, mostly to the west and south
of the city, and significant shrinking of the urban core population (Miamaitijiang et al.,
2014). This shrinkage has led to significant abandonment of properties in the city, resulting
in many vacant lots and decaying infrastructure, mostly between Interstate 44 to the south
and Interstate 270 to the north (Figure 1) (Miamaitijiang et al., 2014; Ganning and Tighe,
2015).
Materials and Methods
Study Area
The city of St. Louis, MO, rests on the western banks of the Mississippi river near
the center of the river valley (Fig. 1). The city has an estimated population of 320,000
people in an area of 170 km2(Ganning and Tighe, 2015). The climate of the city is
considered transitional between humid continental (K¨
oppen climate classification Dfa)
and humid subtropical (Cfa). There is no significant topographical relief in the area,
with the major bodies of water being the Missouri and Mississippi rivers (Fig. 1).
This lack of major geographical features results in the city experiencing very hot and
VOLUME 90, ISSUE 3 177
Fig. 1. Map of the St. Louis, MO, metropolitan area. The demarcations of the political boundaries of the city are
highlighted. For the purpose of this study we defined “city” as the area within the Interstate Highways 255–270 to
the north, west and south, and the Mississippi River to the east. Other shaded areas represent city or county parks.
humid summers (average summer temperature 31.2C, record high 46.1C) and very cold
winters (average winter temperature 0.1C, record 22C). The average seasonal snowfall
is 45 cm.
The vegetation of the city of St. Louis is considered to be transitional between oak-
hickory dominated forests and tall grass prairie (Nigh and Schroeder, 2002). Typical of
urban environments, there are many invasive and weedy plant species that tend to dominate
abandoned and vacant lots, as well as riparian habitats and parks (M¨
uhlenbach, 1979). Like
many other cities (Colasanti et al., 2013), there is also a tremendous resurgence and interest
in agriculture across the city. This has resulted in the establishment of many community
gardens and urban farms over the last decade.
For the purpose of this study, we restricted the definition of the city of St. Louis to
the actual city and the surrounding suburbs contained within the Interstate Highway 270
loop to the north and to the west; Interstate Highway 255 to the south; and the Mississippi
River to the east (Fig. 1). This area is almost continuously urbanized from downtown to the
Interstate Highway 255/270 loop.
178 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
Specimen Sources
The bulk of the specimens (>7700) were collected between May 2013 and October 2016
as part of a wildlife conservation survey for the Missouri Department of Conservation.
Most of those collections were done in urban gardens, urban farms, native vegetation sites,
and restoration prairies. Other habitats sampled less often were abandoned and vacant
lots, private gardens, and city parks. Sampling was done via aerial netting. All specimens
from the survey are in the insect collection at Saint Louis University (SLU). Two other
surveys provided significant numbers of specimens. First is the biennial BioBlitz survey of
Forest Park, which started in 2004. These specimens reside in the personal collection of M.
Arduser. This is the largest public park in St. Louis with an area of 523.25 ha, and it contains
two restoration prairie sites, a savannah-type area, and a continuously forested area. The
second survey was that of pollinators of the Litzsinger Road Ecology Center, conducted by
R. Clinebell and M. W. Slagle in the late 1990’s. This is an educational center managed by
the Missouri Botanical Garden. It is located in the suburbs just west of the city, with an area
of 13.75 ha, and contains a range of natural habitats. Specimens of this survey are deposited
in the collection of the Missouri Botanical Garden (MBG) and the Litzsinger Road Ecology
Center (Litzsinger). Other sources of material were specimens from the collections at The
Enns Entomology Museum, University of Missouri, Columbia (UMC); The University of
Missouri, St. Louis (UMSL); and the Saint Louis Zoo (SLZ). Finally, an amateur collector,
Mr. George Diehl, also contributed specimens. All specimens were identified to species or
morphospecies level. We followed the taxonomical convention of Michener (2000) as per
modifications by Ascher and Pickering (2016). Noticeable in the UMC collection were a
number of specimens collected by the US Department of Agriculture in the late 1930’s as
part of a survey of the invasive Japanese beetle, Popillia japonica, in St. Louis. According
to a report by M. E. Brown in 1996, most of the trapping in the 1930’s St. Louis occurred
between what is now the Gateway Arch grounds to the east, and Tower Grove Park to the
west. This park is adjacent to the grounds of the Missouri Botanical Garden.
A word of caution concerning methods is relevant here. Specimens for this study were
collected using a broad range of methods at different times, e.g., aerial netting in the 2010’s
and flight intercept trap in the 1930’s. For many individuals we don’t have any idea how they
were collected or the specific locality within the city. Specimens were also collected from
the early 20th century until 2016. That’s almost 100 years difference. Thus, determining
various diversity parameters is not appropriate in this case.
Results
A total of five families, 47 genera, and 198 species of bee species were identified as
occurring in the St. Louis metro area (Table 1). The bulk of the species richness found
in the city are native bees (189/198). The most species rich group was the family Apidae
with 20 genera and 61 species. The family of the leaf-cuttter bees, Megachilidae, was the
next most species rich with 12 genera and 45 species. The sweat bees,Family Halictidae,
contained 10 genera and 44 species, whereas the mining bee family, Andrenidae, was
represented by five genera and 38 species. The polyester bees, Family Colletidae, were the
least diverse with two genera and 10 species. The only family reported for the state that was
not observed in the city was Melittidae. Fourteen species were identified from collection
specimens and have not been observed in recent surveys (Table 1). Andrena geranii was
reported in the literature by Rau (1934), yet, we have not identified a voucher specimen for
this species.
VOLUME 90, ISSUE 3 179
Bumble Bees
A total of nine species of the genus Bombus were recorded in the city (Table 1). Seven
of the nine currently occur in the city and have been collected repeatedly in the last
four years. Alternatively, one species, the variable cuckoo bumble bee, B. variabilis,is
represented by a single individual collected in 1938 in the USDA Japanese beetle sur-
vey. The host species of the variable cuckoo bumble bee, the American bumble bee,
B. pensylvanicus (Williams et al., 2014), is consistently found in three locations in the
city.
The rusty patched bumble bee, B. affinis, was collected by R. Clinebell in the Litzsinger
Road Ecology Center. This collection consisted of two males and five workers in late
summer of 1998. This species has not been observed in the St. Louis area ever since.
Bombus affinis was listed as endangered by the US Fish & Wildlife Service on March 21,
2017.
Another species of high conservation concern is the southern plains bumble bee, B.
fraternus. This species has exhibited significant declines in abundance and range over the
last decade (Colla et al., 2012; Hatfield et al., 2014), and is listed by the IUCN as endangered
(Hatfield et al., 2014). We have identified two locations within the city that have populations
of this species, one an urban farm, the other a restoration prairie.
The eastern common bumble bee, B. impatiens, was the most prevalent bumble bee in
all agricultural sites. It was also present in all native vegetation sites, and only absent from
vacant lots. This species was active from late May all the way until the middle of October
in all years that we sampled.
Non-natives Bees
A total of nine non-native bees were identified as occurring in the city (Table 1). The most
common non-native across the entire city was the honeybee, Apis mellifera, with as much
as ten times higher abundance than any other species. Even when hives were not observed
in the vicinity, honeybees were prevalent. Over half of the sites surveyed had hives present,
or hives were observed nearby.
The second most common non-native was the European wool carder bee, Anthidium
manicatum. This species was found mostly in community gardens and urban farms. It was
much less abundant, or even absent, from areas that contained mostly native vegetation.
This species was commonly associated with lamb’s ears, Stachys byzantina. Females of
this species visit the lamb’s ears leaves from which they ‘card’ the trichomes for nesting
material (Smith, 1991).
One of the most recent additions to the bee community in St. Louis is the giant Asian
resin bee, Megachile sculpturalis. It is believed that this species was introduced in the
early 1990’s into the east coast, near Baltimore (Batra, 1998). A decade later it was de-
tected in the state of Kansas (Hinojosa-D´
ıaz, 2008). The earliest record of this species
in the city of St. Louis is also from 2008. This species is mostly found in community
gardens and urban farms associated with Russian sage, Perovskia atriplicifolia.Wenever
observed it in areas that contained only native vegetation. This species has also been iden-
tified pollinating kudzu, Pueraria lobata, in the St. Louis area (S. Callen, unpublished
data).
The third most common non-native was the alfalfa leaf-cutter bee, Megachile rotundata.
This species was also more abundant in agricultural habitats, and almost entirely absent
from native vegetation sites.
180 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
Table 1. Checklist of the bee species reported for St. Louis city, MO. The list is assembled alphabetically for
species within genus, and genera within families. For each species we indicate if the species is native or introduced,
and list the collection(s) where the voucher specimens resides. Species in bold font have not been observed in
recent times within city limits, 1990 to present.
Taxon Origin Voucher
ANDRENIDAE
1Andrena accepta Viereck native Arduser/SLZ
2Andrena barbara Bouseman and LaBerge native Arduser
3Andrena brevipalpis Cockerell native Arduser
4Andrena carlini Cockerell native SLZ
5Andrena cressonii Robertson native SLU/SLZ
6Andrena commodaSmith native Arduser
7Andrena crataegi Robertson native UMC
8Andrena erythrogaster (Ashmead) native UMC/Arduser
9Andrena forbesii Robertson native Arduser
10 Andrena geranii (Robertson)*native
11 Andrena helianthi Robertson native SLU
12 Andrena hippotes Robertson native Arduser
13 Andrena ilicis Mitchell native Litzinger
14 Andrena illini Bouseman and LaBerge native SLZ/UMC
15 Andrena illinoensis Robertson native Litzinger
16 Andrena mandibularis Robertson native UMC
17 Andrena imitatrix Cresson native SLU/SLZ
18 Andrena miserabilis Cresson native SLU/SLZ
19 Andrena nasonii Robertson native SLU/SLZ
20 Andrena nuda Robertson native MBG
21 Andrena perplexa Smith native SLZ
22 Andrena personata Robertson native Litzinger
23 Andrena phaceliae Mitchell native Litzinger
24 Andrena polemonii Robertson native Litzinger
25 Andrena robertsonii Dalla Torre native Litzinger
26 Andrena rudbeckiae Robertson native SLU/SLZ
27 Andrena sayi Robertson native Litzinger
28 Andrena simplex Smith native Arduser/SLZ
29 Andrena violae Robertson native SLU/SLZ
30 Andrena wilkella (Kirby) introduced SLU/SLZ
31 Anthemurgus passiflorae Robertson native SLU
32 Calliopsis andreniformis Smith native SLU
33 Protandrena bancrofti Dunning native UMC
34 Protandrena cockerelli Dunning native UMC
35 Pseudopanurgus albitarsis (Cresson) native Arduser
36 Pseudopanurgus compositarum (Robertson) native Arduser/SLZ
37 Pseudopanurgus labrosus (Robertson) native Arduser
38 Pseudopanurgus rudbeckiae (Robertson) native Arduser
APIDAE
39 Anthophora abrupta Say native SLU
40 Anthophora ursina Cresson native UMC/Arduser
41 Anthophora terminalis Cresson native SLZ/MBG
42 Apis mellifera Linnaeus introduced SLU/SLZ
43 Bombus affinis Cresson native MBG
44 Bombus auricomus (Robertson)native SLU/SLZ
45 Bombus bimaculatus Cresson native SLU/SLZ
46 Bombus fervidus (Fabricius)native SLU
47 Bombus fraternus (Smith)native SLU
VOLUME 90, ISSUE 3 181
Table 1. Continued.
Taxon Origin Voucher
48 Bombus griseocollis (DeGeer)native SLU/SLZ
49 Bombus impatiens Cresson native SLU/SLZ
50 Bombus pennsylvanicus (DeGeer)native SLU/SLZ
51 Bombus variabilis (Cresson)native UMC/MBG
52 Cemolobus ipomoeae (Robertson) native UMC
53 Ceratina calcarata Robertson native SLU/SLZ
54 Ceratina dupla Say native SLZ
55 Ceratina strenua Smith native SLU/SLZ
56 Diadasia australis (Cresson) native SLZ
57 Eucera hamata (Bradley) native SLU/SLZ
58 Eucera rosae (Robertson) native SLU
59 Epeolus bifasciatus Cresson native SLU
60 Florilegus condignus (Cresson) native SLZ
61 Habropoda laboriosa (Fabricius) native SLU/SLZ
62 Holcopasites calliopsidis (Linsley) native SLU
63 Melissodes agilis Cresson native SLU/SLZ
64 Melissodes bimaculatus Lepeletier native SLU/SLZ
65 Melissodes boltoniae Robertson native Arduser
66 Melissodes communis Cresson native SLU
67 Melissodes comptoides Robertson native SLU/SLZ
68 Melissodes coreopsis Robertson native Arduser
69 Melissodes denticulatus Smith native SLU/SLZ
70 Melissodes dentiventris Smith native Arduser/SLZ
71 Melissodes desponsus Smith native SLU/SLZ
72 Melissodes druriellus (Kirby) native Arduser/SLZ
73 Melissodes trinodis Robertson native SLU/SLZ
74 Melissodes vernoniae Robertson native SLU/UMC
75 Melitoma taurea (Say) native SLU/SLZ
76 Nomada texana native SLZ
77 Nomada sp. 1 native SLU
78 Nomada sp. 2 native SLU
79 Nomada sp. 3 native SLU
80 Nomada sp. 4 native SLU
81 Nomada sp. 5 native SLU
82 Nomada sp. 6 native SLU
83 Nomada sp. 7 native SLU
84 Nomada sp. 8 native SLU
85 Nomada sp. 9 native SLU
86 Peponapis pruinosa (Say) native SLU/SLZ
87 Ptilothrix bombiformis (Cresson) native SLU/SLZ
88 Triepeolus atripes Mitchell native Arduser
89 Triepeolus concavus (Cresson) native SLU
90 Triepeolus helianthi (Robertson) native Litzinger
91 Triepeolus lunatus (Say) native SLU
92 Triepeolus quadrifasciatus (Say) native SLU
93 Triepeolus remigatus (Fabricius) native SLU
94 Triepeolus simplex Robertson native Litzinger
95 Triepeolus sp. native SLU
96 Svastra obliqua (Say) native SLU/SLZ
97 Xenoglossa strenua (Cresson)native SLU
98 Xenoglossa kansensis Cockerell native SLZ
99 Xylocopa virginica (Linnaeus) native SLU/SLZ
182 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
Table 1. Continued.
Taxon Origin Voucher
COLLETIDAE
100 Colletes compactus Cresson native Arduser
101 Colletes inaequalis Say native SLU
102 Colletes latitarsis Robertson native UMC
103 Hylaeus affinis (Smith) native Litzinger
104 Hylaeus fedorica (Cockerell) native SLU
105 Hylaeus illinoisensis (Robertson) native SLU/SLZ
106 Hylaeus leptocephalus (Morawitz) introduced SLU
107 Hylaeus mesillae (Cockerell) native SLU/SLZ
108 Hylaeus modestus Say native SLU/SLZ
109 Hylaeus sp. Litzinger
HALICTIDAE
110 Agapostemon virescens (Fabricius) native SLU/SLZ
111 Agapostemon sericeus (Forster) native SLZ
112 Agapostemon splendens (Lepeletier) native SLU
113 Agapostemon texanus Cresson native SLU
114 Augochlora pura (Say)native SLU/SLZ
115 Augochlorella aurata (Smith) native SLU/SLZ
116 Augochlorella persimilis (Viereck) native SLU
117 Augochloropsis fulgida (Smith) native SLU/SLZ
118 Augochloropsis metallica (Fabricius) native UMC
119 Dieunomia heteropoda (Say) native SLU
120 Dieunomia triangulifera (Vachal) native UMC
121 Dufourea marginata (Cresson) native UMC
122 Halictus confusus Smith native SLU/SLZ
123 Halictus ligatus Say native SLU/SLZ
124 Halictus parallelus Say native UMC/Arduser
125 Halictus rubicundus (Christ) native SLU/SLZ
126 Lasioglossum bruneri (Crawford) native SLZ
127 Lasioglossum callidum (Sandhouse) native SLZ
128 Lasioglossum cattellae (Ellis) native Arduser
129 Lasioglossum cinctipes (Provancher) native SLU
130 Lasioglossum coeruleum (Robertson)native SLU/SLZ
131 Lasioglossum coriaceum (Smith) native Arduser
132 Lasioglossum cressonii (Robertson)native SLU/SLZ
133 Lasioglossum ephialtum Gibbs native SLU
134 Lasioglossum forbesii (Robertson) native SLZ
135 Lasioglossum foxii (Robertson)native SLZ
136 Lasioglossum hitchensi Gibbs native SLU/SLZ
137 Lasioglossum illinoense (Robertson) native SLZ
138 Lasioglossum imitatum (Smith)native SLU/SLZ
139 Lasioglossum lustrans (Cockerell) native SLU
140 Lasioglossum obscurum (Robertson)native SLZ
141 Lasioglossum oenotherae (Stevens)native UMC
142 Lasioglossum pectinatum (Robertson) native SLU
143 Lasioglossum pectorale (Smith) native SLU/SLZ
144 Lasioglossum pictum (Crawford) native Litzinger
145 Lasioglossum pilosum (Smith) native SLU
146 Lasioglossum platyparium (Robertson) native Arduser
147 Lasioglossum tegulare (Robertson) native SLU/SLZ
148 Lasioglossum truncatum (Robertson) native Litzinger
VOLUME 90, ISSUE 3 183
Table 1. Continued.
Taxon Origin Voucher
149 Lasioglossum versatum (Robertson) native Arduser
150 Lasioglossum zephyrum (Smith)native SLU/SLZ
151 Nomia nortoni Cresson native UMC
152 Sphecodes heraclei Robertson native Diehl
153 Sphecodes sp. native SLU
MEGACHILIDAE
154 Anthidiellum notatum (Latreille) native SLU
155 Anthidium manicatum (Linnaeus) introduced SLU/SLZ
156 Anthidium oblongatum (Illiger) introduced SLU/SLZ
157 Anthidium psoraleae Robertson native SLU
158 Chelostoma philadelphi (Robertson) native SLU
159 Coelioxys germanus Cresson native UMC
160 Coelioxys hunteri Crawford native UMC
161 Coelioxys modestus Smith native MBG/UMC
162 Coelioxys octodentatus Say native SLU
163 Coelioxys obtusiventris Crawford native SLU
164 Coelioxys sayi Robertson native SLU
165 Dianthidium curvatum (Smith) native SLZ
166 Heriades carinata Cresson native SLU
167 Heriades leavitti Crawford native Arduser/SLZ
168 Heriades variolosa (Cresson) native Arduser
169 Hoplitis pilosifrons (Cresson)native SLU/SLZ
170 Hoplitis producta (Cresson) native SLU
171 Megachile addenda Cresson native SLU
172 Megachile apicalis Spinola introduced SLU
173 Megachile brevis Say native SLU/SLZ
174 Megachile campanulae (Robertson)native SLU/SLZ
175 Megachile concinna Smith introduced SLU/SLZ
176 Megachile exilis Cresson native SLU/SLZ
177 Megachile frugalis Cresson native SLU
178 Megachile gemula Cresson native Litzinger
179 Megachile inimica Cresson native SLU/SLZ
180 Megachile latimanus Say native UMC
181 Megachile mendica Cresson native SLU/SLZ
182 Megachile montivaga Cresson native SLZ
183 Megachile parallela Smith native UMC
184 Megachile petulans Cresson native Arduser
185 Megachile policaris Say native UMC/Arduser
186 Megachile texana Cresson native SLU/SLZ
187 Megachile rotundata (Fabricius) introduced SLU/SLZ
188 Megachile sculpturalis Smith introduced SLU/SLZ
189 Megachile xylocopoides Smith native SLU/SLZ
190 Osmia atriventris Cresson native UMC
191 Osmia bucephala Cresson native Arduser
192 Osmia cordata Robertson native UMC
193 Osmia georgica Cresson native SLU
194 Osmia lignaria Say native SLU
195 Osmia pumila Cresson native SLU/SLZ
196 Osmia subfasciata Cresson native SLU
197 Stelis louisae Cockerell native Arduser
198 Stelis lateralis Cresson native SLZ
*This species is only known from St. Louis by a report in the scientific literature (Rau, 1934), and no voucher specimen
currently exists.
184 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
Cleptoparasites
We recorded a total of 32 species of cleptoparasitic bees in the city of St. Louis. That is
16% of the bee fauna, a higher percentage than what has been reported for New York City
(Matteson et al., 2008), Chicago, IL (Pearson, 1933; Tonietto et al., 2011; Molumby and
Przybylowicz, 2012), and even a restoration prairie located 135 miles north of St. Louis
(Geroff et al., 2014).
Of notable interest is the presence of Ceolioxys obtusiventris. This species is one of the
most rare bees of North America, with only a handful of females (<20) ever found (Ascher
and Pickering, 2016). This species was originally described based on a single specimen
from Florida (Crawford, 1914), and then a second specimen reported in Indiana (Chandler,
1969). We have collected this species twice in an urban farm in the suburb of Ferguson,
with both occasions taking place in the month of July 2014 and 2016.
Discussion
We identified 198 bee species in the city of St. Louis. In the state of Missouri, there are
six families and 452 reported bee species (M. Arduser, in preparation). Almot 45% of the
bee fauna of the state of Missouri has been recorded in the city of St. Louis (198/452). In
terms of raw species richness, St. Louis bee diversity is higher than that of restored prairie
systems in the Midwest United States (Geroff et al., 2014), and is comparable to the natural
environment that is the Indiana Dunes (Grundel et al., 2011).
In general, the flora and fauna of cities tend to be dominated by generalists, with a
significant amount of non-native species (McKinney, 2008). We observed a relatively low
number of introduced species (Table 1), especially when compared to studies from Chicago
(Tonietto et al., 2011; Molumby and Przybylowicz, 2012) and New York (Matteson et al.,
2008). There are two general possibilities for this. First, being located in the center of the
continent provides a geographic barrier to non-natives, given that most of those species
arrive at ports of entry in coastal cities (Lockwood et al., 2013). The second possibility is
that the dominance of the native bee species can keep some of the non-native species from
establishing (Lockwood et al., 2013). These two hypotheses are not mutually exclusive,
and could be effectively interacting in the St. Louis environment.
We did record several specialist bees across the city. The hibiscus bee, Ptilothrix
bombiformis, was recorded from community gardens that featured rose mallow, Hibis-
cus moscheutos, or Rose of Sharon, Hibiscus syriacus.Ptilothrix was collected mostly in
these plants, but in one occasion was collected from a cultivated variety of iris. We recorded
three species of squash bee: Peponapis pruinosa, Xenoglossa kansensis, and X. strenua.We
also collected several specialists on sunflowers, like Svastra obliqua and Dieunomia het-
eropoda. Furthermore, all cleptoparasitic bees are essentially specialists. Within the city of
St. Louis, we identified 32 species of cleptoparasitic bees (Table 1). This group is proposed
to be indicators of the overall stability of the bee community given that cleptoparasites
are host specialists and require the presence and abundance of the host species in order to
maintain a viable population (Scheffield et al., 2013).
There is paucity in the reporting of native bee diversity for most habitats. A survey
of the bees of the Indiana Dunes, a botanical rich native habitat in northwestern Indiana,
revealed a total of 175 species (Grundel et al., 2011). The most species rich genus in the
Indiana Dunes habitat was Lassioglossum, with 25% of the species. In our study, although
the genus Lassioglossum was also the most species rich (20 spp.), yet, it contained only
12% of the species (Table 1). This is likely due to the fact that many Lassioglossum species
VOLUME 90, ISSUE 3 185
are sandy soil specialists and the dunes provide an ideal habitat. The Indiana Dunes, which
represents less than 0.1% of the area of the state, contain nearly half of the bee species
in the state. While this is likely the result of a combination of biogeographic, edaphic and
climatic forces (Grundel et al., 2011), the high bee diversity in the city of St. Louis is likely
the result of socioeconomic and ethnic processes.
For cities, there is an even greater lack of reported bee diversity. The state of Illinois
contains some 500 species of bees (R. Tonietto, personal communication). Pearson (1933)
reported 169 for the Chicago region. Yet, many of the localities that he lists are at con-
siderable distances from what even today are urbanized areas. A more recent estimate of
the bee diversity for the greater Chicago metro area, which is the third largest city in the
United States and more than twice the area of St. Louis, is 93 species (Molumby and
Przybylowicz, 2012). Therefore, the Chicago area contains less than one fifth of the state’s
bee fauna. Furthermore, the bee diversity within the city limits of is estimated to be 68
species (Minor et al., 2016). The large bee diversity in the city of St. Louis is noteworthy
and must be further investigated to determine the specific characteristics that maintain this
large diversity.
Over the last decade there has been increased concern over the conservation status of
many pollinators at the national (National Academy of Sciences, 2007; USA President’s
Task Force Strategy on Pollinator Health, 2015) and international levels (IPBES, 2016).
Many food producing systems, and thus food security, depend on pollinator services mostly
provided by bees. Furthermore, many wildlife species depend also on pollination services
for their foraging and nutritional needs. A recent report by the Intergovernmental Science-
Policy Platform on Biodiversity and Ecosystem Services (IPBES, 2016) estimates that as
many as 40% species of pollinators worldwide are declining, threatened, or endangered. For
many conservation practitioners involved in managing pollinator populations, it is crucial
to understand how much diversity is present and how is distributed. Traditionally, efforts
to conserve pollinator biodiveristy have been mostly focused on natural areas that could
be protected or managed with relatively minimal human intervention. We need to consider
that at times part of the pollinator conservation strategy might be “the city.” There is no
denying that urbanization has resulted in significant loss of biodiversity (McKinney 2008,
Butchart et al., 2010), and pollinators are not immune (Potts et al., 2010, Hadley and Betts,
2012). Yet, recent studies (Baldock et al., 2015; Ives et al., 2015; Hall et al., 2017) suggest
that we need to incorporate the role that novel urban ecosystems have in our understanding
for the biological conservation of pollinators.
Acknowledgments
We wish to acknowledge Robert Sites (UMC), Peter Hoch (MBG) and Robert Marquis
(UMSL) for access to the respective collections and the loaning of specimens. We also wish
to thank Malinda Walter Slagle and George Diehl for access to specimens and information
regarding the original survey of the Litzsinger Road Ecology Center. Thanks to Erin Shank
(Missouri Department of Conservation) for assistance with access to sites. Rebecca Toni-
etto, Peter Bernhardt, Damon Hall, and Lara Zwarun provided valuable feedback on this
manuscript. Special thanks to the dozens of undergraduate students in GRC’s lab that have
assisted over the years with collecting, pinning, labeling, sorting and cataloging of speci-
mens. Funding for this study was provided by the Missouri Department of Conservation’s
Wildlife Diversity Fund, Saint Louis University’ Center for Sustainability Innovation Fund,
and the President’s Research Fund.
186 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
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... Large tracts of the USA, such as the High Plains and Great Plains are poorly sampled, even though they may have some of the greatest habitat loss across the country (Samson et al., 2004). With limited sampling effort, diverse bee communities can be found in both pristine (e.g., Grundel et al., 2011) and in anthropogenically disturbed habitats (e.g., Camilo et al., 2017). For example, a single day of collecting in Bellaire, Michigan added 50 new bee species records to Antrim County, bringing the total known species from 34 to 84 in 2016 (Gibbs unpublished data). ...
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