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INTRODUCTION
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Figure 1. Outline of experiments conducted between the years 1996 and 2015, bee preference studies (A) and pollination efficacy and
foraging studies (B). Cage studies are surrounded by dashed line boxes and open field studies are surrounded by solid line boxes.
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
030
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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RESULTS
Study
Bee Preference Studies
Blueberry Flower Selection Choice By Bumble
Bees, Flower Size And Age - Over both years (2007 and
2008) wild blueberry flower corolla length and diameter
were highly correlated (r = +0.527, n=90, P< 0.0001).
Flower age was not correlated with either corolla length
or diameter (P> 0.05). Because of this only year, corolla
length and flower age were modeled to determine if
bumble bees select flowers to visit in choice bioassays.
Bumble bees appear to demonstrate preference for
2
longer wild blueberry flowers (x (1) = 7.221, P = 0.007,
Figure 2), but they did not discriminate for flower age (P>
0.05) when the choice was made between flowers that
differed in age up to 6 days. However, a non-significant
trend was found, which suggests a tendency to select
younger flowers (P = 0.106). Year and the interaction
between corolla length and flower age was also not
significant. The odds ratio for the significant corolla length
effect suggests that for every difference in 1.0 mm length
between flowers that it is 2.3 times more likely that the
larger flower will be first selected by a bumble bee
forager.
Blueberry Flower Selection Choice By Bumble
Bees, Previously Visited Flowers - The trials that focused
on previously visited flowers compared to flowers that
had never been by a bumble bee showed no evidence of
choice (P = 0.715) and the trials assessing visitation to
flowers that had been previously visited from 1 to 6 times
showed no evidence of an effect on bumble bee foragers
(P = 0.363).
Recruitment
Of
Foraging
Honey
Bees,
Commercial Bumble Bees And Native Wild bees To
Clone Flower Density In The Field - I found that floral
density per stem (both open and non-opened flowers) in
clones did determine the recruitment of honey bees,
bumble bee queens, and native bees other than bumble
bees, but this effect was dependent upon site (Table
1).With regards to the entire foraging bee community (all
bee taxa), bees recruited to clones with higher total
flower density. There was an effect of bee density due to
site which is not unexpected as bee densities, especially
native bees, vary by an order of magnitude from field to
field and geographic region to region.
In the field,the frequency at which stems were
selected by foraging bumble bees in Jonesboro, Maine
was not influenced by clone (genet; P = 0.178), nor the
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Figure 2. Modeled probability of a bumble bee forager selecting a flower when given a choice of two flowers. As number become
more negative on x-axis the flower chosen was smaller compared to larger flower (mm), as number become more positive, the flower
chosen was larger compared to smaller flower (mm).
Significant Factors
F-test
site
flower density
site
flower density
flower density
site
flower density
site x flower density
Proportion
variance
2
explained, r
0.584* (0.418)**
0.331 (0.224)
0.364
0.512 (0.350)
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Figure 3. Preference of Bombus impatiens workers (stipled bar) for species other than blueberry (measured in paired bioassays as
the percentage of first landings on blueberry vs the other species). In addition the relative time spent on blueberry flowers relative to
the co-flower (hatched bar). The red dashed line indicates a region where foragers spent the same time on blueberry and the coflowering plant and the blue dotted line indicates the region of no-preference in initial floral choice between blueberry and a coflowering plant species. Asterisks indicate significant differences in 1) time spent on the co-flower relative to blueberry and 2)
preference for choice of initial flower selection.
Foraging Studies
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Figure 4. Average pollen deposition on floral stigmas after a single visit and flower handling time by common wild blueberry pollinators: bumble bees,
B. impatiens, digger bees, Andrena carlini, leaf cutting bees, O. atriventris, and honey bees, A. mellifera. Bars with the same letters (uppercase
handling time, lowercase pollen deposition) are not significantly different, Tukeys test, experiment-wise error rate at P< 0.05.
Figure 5. Pollen deposition on a floral stigma by a bumble bee visiting a flower previously visited by a bumble
bee and then visiting a new flower (BB/BB), or by a bumble bee visiting a flower previously visited by a honey
bee and then visiting a new flower (HB/BB), or by a honey bee visiting a flower previously visited by a honey
bee and then visiting a new flower, or by a honey bee visiting a flower previously visited by a bumble bee and
then visiting a new flower (BB/HB). Bars with the same letters are not significantly different, Tukeys test,
experiment-wise error rate at P< 0.05.
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Figure 6. Diurnal foraging periodicity of honey bees and native bees, 20-31 May, 2001 in
Winterport, Maine.
Figure 7.Relationship between air temperature at time of sampling and the percent of a specific taxon groups population (native solitary
bees, bumble bee queens, and honey bees) for a given day.
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Table 2. Probability density function fit to the frequency distributions of the number of stems per bout, distance between
stems (m), cardinal direction to the next stem, and flowers visited per stem for four taxa of common bee pollinators of wild
blueberry in Maine.
bee taxon
group
sample size
of measure
measure
distribution
parameters
goodness of
fit*
Osmia
spp.
47
stems / bout
38
87
Gamma
Poisson
Poisson
15
stems / bout
Exponential
0.118 (P)
45
Gamma
Poisson
Gamma
25
Weibull
81
173
2508
Weibull
2508
2536
u = 2.767
= 1.061
na**
0.779 (LR)
24
610
stems / bout
Beta
4509
Exponential
Andrena
spp.
Bombus
spp.
Honey
bee, A.
mellifera
4903
5333
Gamma
Poisson
Exponential
Log Normal
uniform
Beta
uniform
Poisson
na
0.073 (CM)
0.708 (P)
0.351 (P)
0.150 (K)
0.364 (K)
0.999 (LR)
0.10 (D)
0.994 (LR)
na
0.204 (K)
0.251 (K)
0.298 (LR)
na
0.999 (P)
* probability value for goodness of fit to theoretical distribution, (test statistic) = P: Pearson chisquare, CM: Cramer von Mises, K:
Kolmogorovs D, LR: adequacy LR test.
** na: not applicable.
DISCUSSION
Over 120 species of bees have been found to be
associated with wild blueberry in Maine (Bushmann and
Drummond, 2015). Little is known, however, about their
behavior and activity during the blooming season. The
collection of studies, reported here, begin to shed light on
how individual bee species or groups of species play a
role in wild blueberry pollination.
There are several major areas that I would like to address
in this discussion.
First of all, it is apparent that bees, as shown by
our work with bumble bees, make choices both at the
blueberry flower level, but also at the stem floral density
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Figure 8. Frequency distribution for distance between stems moved by honey bees (a) and Andrena spp.
(b).
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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REFERENCES
CONCLUSION
An understanding of crop pollination requires an
understanding of the interactions between: bees and crop
plant reproductive biology, as well as among morphology,
bee species, the abiotic environment and bee behavior
and plant behavior and physiology. This work reports on
several isolated studies that begin to increase our
understanding of several of these interactions. The
contribution of these studies is in providing quantitative
relationships that can be used to construct a computer
simulation model of bee pollination of wild blueberry.
Specifically, I have shown that bumble bees
discriminate and select larger flowers, but that flower age
or whether flowers were previously visited did not affect
flower selection. Flower density does affect long-distance
recruitment to clones, with clones characterized by higher
flower density receiving higher bee visitation. I also found
that blueberry flowers are much more attractive than five
other native co-flowering shrubs. This is important
because highly attractive shrubs that flower at the same
time as blueberry might pull bees away from the
blueberry crop during pollination. Another important
finding is that native bees are much more efficient, on a
per bee basis, than the introduced honey bee at
pollinating blueberry, in terms of flower handling time and
pollen deposition on the stigma of the flower. However,
the presence of bumble bees visiting flowers prior to
honey bees appears to increase the pollination efficacy of
honey bees. The last important finding in this study that
has direct implications to modeling pollination is that I
found differential lengths of time during the day that
different taxa foraged on blueberry and that this was
partly explained by air temperature. Bee movement in
blueberry fields was found to be dependent upon the
taxon. This aspect of bee foraging has not been
investigated in simulation models constructed to
investigate the dynamics of crop pollination. I believe that
all of my findings will allow the construction of a
preliminary model.
ACKNOWLEDGEMENTS
I would like to thank the assistance of Constance Stubbs,
Stephanie L. Allard, Lisa Campbell, Judith Collins, and
several undergraduate research assistants that helped on
data collection for these studies over the past 20 years.
This is Maine Agricultural and Forest Experiment Station
Publication3500.Financial support for some of the
objectives in this project (2011-2015) was provided by the
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
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Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA
Drummond FA.
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Stubbs CS, Jacobson HA, Osgood EA, Drummond FA
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Stubbs CS, Drummond FA (2001).Bombus impatiens
(Hymenoptera: Apidae): An alternative to Apis mellifera
Copyright: 2016 Drummond FA. This is an openaccess article distributed under the terms of the Creative
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use, distribution, and reproduction in any medium,
provided the original author and source are cited.
Behavior of Bees Associated with the Wild Blueberry Agro-ecosystem in the USA