Table Of ContentPROC. ENTOMOL. SOC. WASH.
109(1), 2007, pp. 249-252
Note
Biological Notes on Heriades carmatus Cresson, Heriades leavitti Crawford,
and Heriades variolosus (Cresson) (Hymenoptera: Megachilidae)
Biological information is available for a Bullrun Creek at the edge of a pasture
three of the 13 North American species on Fulker Farm; 5) 4 km south of
ofthe bee genus Heriades Spinola (Mega- Bozeman along Hyalite Creek; 6) 3 km
chilidae: Osmiini) (Griswold 1985): Her- east of Bozeman between a stream and
iades carinatus Cresson (Rau 1922; a cultivated field on Rocky Creek Farm;
Matthews 1965; Krombein 1967; Jensen and 7) Bozeman at the Montana State
et al. 2003), Heriades leavitti Crawford University Horticultural Farm. Most H.
mm
(Krombein 1967), and Heriades variolo- carinatus nested in 3.2 and 3.7
sus (Cresson) (Fischer 1955). Here, we diameter tunnels. Of the 254 emerging
describe the structure of H. carinatus H. carinatus adults that we could asso-
nests in Montana and compare it to ciate with nests of known diameter, 105
mm
observations from other locations. For came from 3.2 nests, 134 from
mm mm
all three species, we add data on emer- 3.7 nests, ten from 4.6 nests,
mm
gence sex ratios, and for H. leavitti, we three from 5.9 nests, and one each
mm
report records for nest associates, in- from 7.5 and 8.0 nests. Matthews
cluding Sapyga spp. (Sapygidae). (1965) found that H. carinatus usually
mm
To construct trap-nests, we drilled nested in 3.1 holes in Michigan and
15 cm long holes of varying diameter 3.5 mm holes in Oregon, differences
into pine boards and then inserted paper between the sites being attributable to
tubes with internal diameters of 3.2, 3.7, the different sizes of trap-nests placed in
4.6, 5.9, 7.5, 8.0, and 9.0 mm. Nest the two locations. When Krombein
boards, with the nest holes facing south- (1967) provided trap-nests ranging from
mm
east, were placed adjacent to trees at 3.2-12.7 in diameter, H. carinatus
m
heights of 1.5-2.0 at sites in Gallatin used only the two smallest hole sizes, 3.2
Co., Montana, during the last week of and 4.8 mm. Thus, the choice of nest
May 1999 and 2000 (by PDJ and KMO) hole sizes appears to be relatively con-
and Seneca Co., New York, on 16 May stant among populations.
2002 (by KMO and JFO). We collected Of the ten nests dissected, seven
completed nests and held them at room contained only H. carinatus cells, where-
temperature until late fall each year when as three also contained cells of Trypox-
they were transferred to a 8°C /85% ylon (Trypoxylon) sp. (Hymenoptera:
relative humidity cold room. The follow- Crabronidae). The latter included one
ing April, nests were returned to room nest with two H. carinatus cells outside
temperature for post-di—apause rearing. offive Trypoxylon cells, one with four H.
Heriades carinatus. We found H. carinatus cells outside oftwo Trypoxylon
carinatus in trap nests at seven sites in cells, and one with two Trypoxylon cells
Gallatin Co., Montana: 1) 13 km S. of outside of one H. carinatus cell.
Three Forks along the Madison River; The inner portion of seven completed
2) 5.2 km north of Logan along an Heriades-on\y nests included a basal
intermittent stream; 3) 3 and 5 km space (0-69 mm long; mean ± SE =
northeast of Norris along a stream; 19.3 ± 9.8). Next, was a sequence of2-9
4) 5 km northeast of Manhattan along contiguous brood cells (mean = 5.4 ±
250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1.1 per nest), 6-13 mm long (mean = 8.8 0.001). The 15 bees that emerged from
± 3 mm; N = 36). Each cell was 4.6-8.0 mm nests included nine females
delineated from others and from the (X2 = 0.60, P > 0.50). No one site
basal space by 1-2 mm thick partitions produced less than 74% females. Anoth-
of plant resin, but cells were not lined er group of nests, for which we did not
with any visible material. The fmal cell in record the diameter of tunnels from
each nest was followed by one (N = 4) or which each bee emerged, produced
two empty "vestibular" spaces. When 83.9% females (N = 155, x^ = 71.1, P
two vestibular spaces were present, they < 0.001). The strong and consistent bias
were separated from each other by a 1- towards females contrasts with Mat-
mm
3 thick resin plug. The total vestib- thews (1965) results from Michigan
ular space, including that in nests in where he observed emergence sex ratios
which Heriades superseded Trypoxylon, of 26% females in 1963 (N = 57, x- =
ranged from 22-122 mm (mean = 66.3 12.8, P < 0.001) and 71% females in
± 12.0 mm). Each of the nine nests 1964 (N = 21, X- = 3.86, P < 0.05) (our
finished by a Heriades female was analysis of his data). In trap-nesting
capped with a final 3-7 mm thick resin species, it is common for cells constructed
plug (mean = 4.8 ± 0.5 mm). Overall, fordaughters(innercells)tobelargerthan
34.0 ± 7.5% of space in Heriades-only those made for sons (Krombein 1967).
nests was devoted to brood cells and However, we found no correlation be-
their partitions, with empty space com- tween cell position and cell length (Spear-
prising 62.5 ± 7.8% and the final plug mann rank correlation, P = 0.82; N = 36)
3.5 ± 0.4%. Matthews (1965), who and no difference between the length of
provided shorter nest tunnels, reported the innermost and outermost cell in each
no basal spaces in H. carinatus nests, so nest (Wilcoxon Signed Rank Test, P =
that the initial cells were either directly 0.09). Similarly, Matthews (1965) found
against the inner end of the boring or no difference in the size of male and
against a resin plug placed at the inner femaleH. carinatuscells, althoughfemales
end. Like Krombein (1967), we found no tended to occupy in—ner cells.
plugs delineating the inner border of Heriades leavitti. The single previous
basal spaces. Matthews and Krombein report of H. leavitti nests was by
both reported vestibular cells, although Krombein (1967), who found five nests
mm
they were absent in 13% of 89 nests at Lake Placid, Florida, all in 3.2
examined by Matthews. The range in the diameter tunnels; he reared just one
number of cells in Montana nests was individual of each sex from nests, and
nearly identical to that seen by Matthews found no nest associates. In 2002, we
mm
(1965) in Michigan. Because nest holes found H. leavitti in seven 3.2 nests
were shorter in Michigan, the similar along the south edge of the North Pool
number is likely due to lack of a basal at Montezuma National Wildlife Refuge,
space in Michigan nests. The greater Seneca Co., New York. This extends our
number of cells Matthews observed in knowledge of the distribution of H.
Oregon (mean = 6.4, range: 2 13), was leavitti, which had not been previously
offset by shorter vestibular cells. reported in New York State (Hurd 1979;
Sex ratios of emerging H. carinatus J. Ascher, personal communication). The
from all Montana sites combined were nests produced 25 adult female and 18
highly skewed towards females in aduh male H. leavitti {j} = 1.14, P >
3.2 mm nests (94.4% female, N = 108; 0.10), but five nests also had other
j} = 85.3, P < 0.001) and 3.7 mm nests occupants (Table 1). The single nest that
(78.1% females, N = 137; x^ = 43.3, P < we dissected contained ten cells whose
) )
VOLUME NUMBER
109, 1 251
Table 1. Contents ofH. leavitti nests from the Montezuma National Wildlife Refuge.
Nest Heriadesleavitti
number OtherNestOccupants
1 5 Hylaeus annulatus (L.)' (5 c?, 3 ?); Sapyga sp.(l (?, 2 ?)
2 1 H. annulatus (2 c?)
3
4 7 Sapyga louisi Krombein (1 S, 2 ?
5 10
6 1 S. louisi (4 2
7 1 H. carinatus (2 ?)
Total 25
1 = Hylaeus ellipticus (Kirby) (Colletidae).
contents, beginning with the innermost Griswold, and Karl Magnacca for iden-
cell, were: one female S. louisi, one tifying bees. John Ascher provided help-
female H. leavitti, four male H. leavitti, ful comments on the manuscript, and
three female S. louisi, and one male H. shared his knowledge of the distribution
leavitti. In North America, Sapyga have of Heriades. Peter Fay, Marguerite
been reported attacking only Megachili- Fulker, Bret and Katherine Olson, Ro-
dae (Krombein 1979), so it seems likely bert Lane, and the Montezuma Wildlife
that they attacked Heriades rather than Refuge allowed use of their properties
Hylaeus in our nests. If so, Sapyga spp. for the study; Tracy Gingrich, the
infested 18% ofthe Heriades cells in NY. Refuge Biologist, provided assistance in
This is higher than the overall rate locating suitable trap-nesting sites.
reported by Matthews (1965), who ob- Voucher specimens of Heriades have
served S. louisi in 0.4% of H. carinatus been placed in the Montana Entomology
cells, but much lower than the maximum Collection, Montana State University.
rate of 74% for Sapyga pumila Cresson The research in Montana was done in
attacking Megachile rotundata (L.) partial fulfillment ofrequirements for an
(Torchio 1972); in the latter case, how- M.S. in Entomology at Montana State
ever, bees cells were at extremely high University by Peter D. Jensen, and was
densities in populations managed to aid supported by the Montana Agricultural
alfalfa pollination. — Experiment Station.
Heriades variolosus. Two nests col-
lected in 1999 from the Fulker Farm in Literature Cited
Montana were occupied by H. variolosus. Fischer, R. L. 1955. A nest ofHeriades variolosus
One male and 1 1 females emerged from (Cress.); (Hymenoptera: Megachilidae). Cana-
mm
a 3.2 nest, and five females emerged dian Entomologist 88: 33-36.
from a 3.7 mm nest. A 5 mm nest Griswold, T. L. 1985. A generic and subgeneric
revision of the Heriades genus-group. Ph.D.
collected in 2000 produced one male H.
variolosus. The only previous report on HurdT,hePs.isD,.,UtJra.h1S9t7a9t.eAUpnoiivdeersai,typ,p.xii1i7-4i-12-202709pp..In
this species was that of Fischer (1955) Krombein, K. V., P. D. Hurd,Jr., D. R. Smith,
who found a single five-celled nest in and B. D. Burks,eds. CatalogofHymenoptera
hollow sumac {Rhus glabra L.) stem with in America North ofMexico. Vol. 2. Apocrita
a bAocrkenodwialmeedtgemrenotfs3.m—mW.e thank Wil- W(Aacsuhlienagtat)o.n, SDmCi.thxsvoin+iapnp. I1n1s9t9i-tu2t2i0o9n. Press,
Jensen, P. D., K. M. O'Neill, and M. Lavin. 2003.
liam Kemp for providing trap-nesting Pollen provision records for three species of
materials, and Richard Miller, Terry solitary bees (Hymenoptera: Megachilidae;
252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Megac/nle. Heriades) in southwestern Mon- Peter D. Jensen, Kevin M. O'Neill,
tana. Proceedings of the Entomological Soci- James F. O'Neill, and Richard S. Miller.
ety ofWashington 105: 196-203. (PDJ) Department ofEntomology, Mon-
Krombein, K. V. 1967. Trap-nesting Wasps and MT
Bees: Life Histories, Nests, and Associates. tana State University, Bozeman,
Smithsonian Press, Washington, D.C. 570 pp. 59717. U.S.A. (present address: Depart-
. 1979. Sapygidae, pp. 1319-1321. In Krom- ment of Entomology, University of
bein, K. v., P. D. Hurd, Jr., D. R. Smith, and Maryland, College Park, MD 20742,
AB.meDr.icBaurkNso,rtehds.ofCaMteaxliocgo.ofVHoylm.e2n.opAtpoecrraitian U.S.A.); (KMO) Department of Land
(Aculeata). Smithsonian Institution Press. Resources and Environmental Sciences,
MT
Washington, DC. xvi + pp. 1199-2209. Montana State University, Bozeman,
Matthews, R. W. 1965. The biology of Heriades 59717, U.S.A. (author to whom reprint
carinata Cresson. Contributions of the Amer-
requests should be submitted) (e-mail:
ican Entomological Institute 1: 1-33.
Rau, P. 1922. Ecological and behavior notes on koneill(S),montana.edu);N(YJFO) 188 Wood-
Missouri insects. Transactions ofthe St. Louis lawn Avenue, Auburn, 13021, U.S.A.;
Academy ofScience 24: 1-71. (RSM) Department of Land Resources
Torchio,P.F. 1972.SapygapwnilaCresson,aparasite and Environmental Sciences, Montana
ofMegachilerotiindata(¥.)(Hymenoptera: Sapy- MT
gidae; Megachilidae). I: Biology and Description State University, Bozeman, 59717,
ofImmatureStages. Melanderia 10: 1-22. U.S.A.
