JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
68(3), 1995, pp. 279-289
Foraging
in Social Wasps: Agelaia Lacks Recruitment
(Hymenoptera: Vespidae)
L. Jeanne
Robert
and Zoology, University
of Entomology
WI 53706, U.S.A.
Madison,
Departments
Department
to Food
of Wisconsin,
James H. Hunt
of Biology, University
of Missouri,
St. Louis, MO 63121, U.S.A.
Malcolm
G. Keeping*
of Durban-Westville,
4000, South Africa
Zoology Department,
University
Durban
Private Bag X54001,
abstract:
Peru
failed
on Agelaia
in Venezuela
in
and A. hamiltoni
multipicta
to carrion. Foraging
of nestmates
ranges of neigh
and foragers occasionally
showed aggression
when
they encountered
Experimental
to demonstrate
tests
recruitment
boring colonies
overlap
one another
on the baits.
vorable
to engage
in scramble
These
with
appear
species
competition
reason for the
and non-congeneric
wasps,
ants, and bees. A possible
scavenging
to evolve
of food location
these wasps
nest-based
is the unfa
communication
size or to the behavior
ratio due either to inadequate
of inter
benefit/cost
colony
specific
competitors
congeners
failure of
at carrion.
to
the ability
ants, and social bees have evolved
Many
species of termites,
an
to food resources. When
recruit nestmates
discover
item
they
solitary foragers
to other foragers in the nest.
cannot handle alone, they communicate
its presence
Some ant and higher termite species lay chemical
trails that provide distance and
direction
about the food resource
and
information
1971; H?lldobler
(Wilson,
in
bees
and
communicate
distance
direction
Wilson,
1990). Honey
(Apis spp.)
et al., 1985) and
formation via the round and waggle dances (Frisch, 1967; Gould
some of the stingless bees (Meliponinae)
to sources of
lay trails of scent marks
nectar or carrion (Lindauer and Kerr,
is
1958; Roubik,
1989). Based on what
known of ants and bees, the existence
of a recruitment mechanism
appears to be
correlated with large colony size and the utilization
of rich, localized,
ephemeral
food resources (Taylor, 1978; Johnson,
and
1983; Beckers et al., 1989; H?lldobler
Wilson,
Some
1990).
species of social wasps achieve colony sizes of 104 or more (Jeanne, 1991),
to those of ants, bees and termites
that recruit to food. Many
also
comparable
on
resources
as
food
such
carrion
rich, ephemeral
forage
(O'Donnell,
1995). Yet
the social wasps
remain the only group of eusocial
insects for which
(Vespidae)
to a food source has not been clearly documented.
nest-based
recruitment
There
is evidence
for weak recruitment
to concentrated
food sources in some vespines.
When
foragers of Vespula germ?nica
(Fab.) and V. vulgaris (L.) were trained to
dishes of scented honey, more nestmates
arrived at dishes with the same scent
*
Mt.
Present
address: Department
of Entomology,
South Africa.
4300,
Edgecombe
for publication
16 December
1994.
Accepted
South African
Sugar Association
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
Experiment
Station,
280
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
et al., 1974). When
than to nearby control dishes scented differently
(Maschwitz
the Japanese hornet, Vespa mandarinia
Smith, raids a colony of honey bees {Apis
mellifera L.) the number of attacking hornets increases rapidly during the slaughter
phase from two or three to dozens,
(Matsuura
suggesting some form of recruitment
in either case, however,
and Yamane,
that distance
1990). There is no evidence
or directional
information
is communicated
within
the nest or that odor trails
are utilized.
a
the Polistinae,
Within
the species that reproduce
by swarms have evolved
to
swarm-mates
nest
of
chemical
communication
recruit
from
the
natal
system
site to a new nest site selected by scouts (Jeanne,
1981, 1991). This mechanism
to recruit to food (Lindauer,
is similar to that used by stingless bees (Meliponinae)
context
could
be
used
in
the
and
of
food
recruitment.
scutellaris
1961)
Polybia
White
is said to have a primitive
recruitment
communication
of
system lacking
In
distance and direction
asserted
that
contrast, Forsyth
1961).
(Lindauer,
(1978)
he tested 10 species in 6 genera {Metapolybia,
Polybia, Parachartergus,
Synoeca,
and found no evidence
of recruitment.
Neither
Lindauer
Brachygastra,
Agelaia)
nor Forsyth, however,
the type of bait or methods
described
they used or provided
data to support their conclusions.
any quantitative
in the wasps and therefore
Because
food recruitment may be poorly developed
difficult to detect, it would be careless to conclude on the basis of the preliminary
studies done so far that social wasps
lack this ability. Yet the question
of recruit
ment
to food among wasps is important for a full understanding
of the conditions
in the social insects. If the
under which
such communication
systems evolved
it raises the question
social wasps
lack recruitment,
of why.
to test the null hy
Here we report the first controlled
experiments
designed
to
not
in
that
does
nest-based
recruitment
food
exist
pothesis
polistine wasps. We
a Neotropical
21
of
chose to work with Agelaia,
of the
genus
species. Members
are
occurrence
candidates
for
the
of
food
recruitment
because
genus
potential
some species form colonies
in excess of 103 adults (106 in A. vicina) (Jeanne, 1991)
on
and because
vertebrate
carrion (O'Donnell,
they forage
1995). We conducted
our experiments
on A. multipicta
and
A.
hamiltoni
Richards
because
(Haliday)
in Brazil
in our study sites. A colony of A. multipicta
collected
of their availability
contained
5500 adults (Araujo,
1940; Richards,
1978); there are no published
records on colony populations
of A. hamiltoni.
Materials
and Methods
Venezuela
October
studies were done on Agelaia multipicta
1993 at Hato
19-23,
a cattle ranch and biological
station in the southern part of the state of
is at the end of the wet season.
The climate
is seasonally
dry; October
Cojedes.
in
The habitat
includes patches of semi-deciduous
tropical forest and savanna
the waterways.
This
forest borders
terspersed with cleared pastureland.
Gallery
are de
is called the llanos (plains); its plant communities
region of Venezuela
x
m
a
area
were
in
100
Sarmiento
done
50
scribed by
(1983). Our experiments
to
trees
of grass and low herbaceous
ranch
under
shade
large
adjacent
vegetation
of the abundance
of foragers of
buildings
(Fig. 1). The area was chosen because
A. multipicta,
which
is the only member
of the genus we found at this site. An
Field
Pinero,
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
VOLUME 68, NUMBER
1.
Fig.
Venezuela.
Positions
Circles
Dishes
buildings.
show position
of
of bait
represent
belonging
(1) control
281
3
dishes
in relation
to sites
indicate
squares
nests,
to separate
replications
dish and (2) experimental
of known
bait
are
dish
dishes,
labelled
on first
colonies
and
with
of Agelaia
hatched
multipicta
rectangles
different
letters.
trial of each
in
represent
Subscripts
of replications
A, B,
andC.
A. multipicta
thornbird
from the nest of a plain-fronted
(Pha
colony collected
1750 adults. Based on comparisons
of forager
cellodomus
rufifrons) contained
in our study area, the latter were much
traffic at this colony with that of colonies
larger.
to test whether
of food
We designed our experiments
communication
explicitly
location occurs at the nest. We baited foragers with
lumps of raw ground beef,
in petri dishes and set on benches
2 cm in diameter,
ap
placed
approximately
m
use
ants.
the
The
of benches
0.5
off
excluded
proximately
ground.
effectively
them
Baits were refreshed every half hour, either by turning them over, breaking
or
to
them
Each
with
of
moist
fresh
expose
surfaces,
apart
pieces.
by replacing
the three replicates consisted
of two trials run at the same times on two successive
one control dish and one experimental
dish (treatments)
days. Each trial employed
m
11-48
The
the
locations
of
for
the
dishes
three
apart.
spaced
replicates were
selected so as to sample different parts of the 50 x 100 m area (Fig. 1). On the
one of the two dishes was arbitrarily
first day of a replication
the
designated
control (location
1), the other experimental
(location 2). On the second day the
dishes were placed in the same locations at the same hour, but their treatments
were switched
to control for location effects. This experimental
design tests for
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
282
Fig.
Circles
2.
Positions
in relation
of bait dishes
nests, squares
represent
to separate
replicates
belonging
are
to sites of known
colonies
bait dishes,
and hatched
rectangles
letters.
labelled with different
indicate
of Agelaia
represent
hamiltoni
buildings.
in Peru.
Dishes
of the location of a specific resource, but does not
any kind of communication
test for a general alerting at the nest to all resources with the same odor.
A trial was begun with
the simultaneous
placing of baits in the control and
at
the
control dishes were captured with
All
dishes.
experimental
foragers arriving
them from recruiting or attracting
forceps and killed in a killing jar to prevent
nestmates.
dishes were carefully captured
arriving at the experimental
Foragers
and re
with paint (DecoColor?
with forceps, marked
opaque paint markers),
leased. The number of new foragers arriving at each dish during each half-hour
interval was recorded. Each of the 6 trials lasted 2 hours.
Peru
were conducted December
80
13-17,
1993, at Explorama
Experiments
Lodge,
km down the Amazon
second
River from Iquitos, Peru, in an area of disturbed
adjacent to tropical wet evergreen forest. Agelaia hamiltoni was
growth vegetation
the most common member
of its genus in this habitat. This wasp nests in cavities
sizes are
in the carton nests of ants of the genus Azteca
1978). Colony
(Richards,
we
to
When
disturbed
be
for
this
but
appear
colony
unreported
large.
species,
adults emerged
several thousand
#253 following
these experiments,
(estimated)
it.
from the nest to defend
3 x 3cm piece of raw beef [replicate
As in Venezuela,
bait (approximately
1]
or fish [replicate 2]) was placed in a petri dish and set on a stool (approximately
of the Venezuela
0.5 m above ground) to exclude ants. Two modifications
protocol
were made. First, in order to control for the possibility
that a general alerting at
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
VOLUME 68, NUMBER
283
3
and
the nest to the odor of the bait may bring recruits to both the experimental
the
the control dishes,
the control was put out on one day, followed
experi
by
at the same time on the following
mental
bait in the same position
day. The
the results. If the experimental
trial were
control was run first to avoid biasing
run first, it would
the
that could artificially
elevate
train a cohort of foragers
numbers
recorded during the control trial on the second day.
to control for the possibility
that killing foragers at the control dish
Second,
and result in a downward
bias of the
the local foraging population
may deplete
numbers
dish on the next day, each new arrival at the
finding the experimental
control dish was aspirated
from the control bait, marked,
held for the duration
In other words, wasps discovering
of the trial, then released.
the control dishes
were not allowed
to return to their nests with food, but they were returned to the
the
prior to the next day's experiment.
Foragers
foraging population
reaching
dish were captured with forceps, marked,
and immediately
released,
experimental
as before. The number of new foragers arriving in each half hour interval at each
to known
dish was recorded. The position
of the dish in each trial in relation
nests of A. hamiltoni
is shown in Fig. 2. Each of the two control and two exper
imental trials lasted 2.5 hours.
Analysis
of variance on square-root-transformed
data was used to test the null
Analysis
new
the
of
and
numbers
that
hypothesis
foragers arriving at the experimental
control baits did not differ (Systat for Windows,
1992).
Results
Venezuela
new arrivals at the experimental
and marking
dishes did not appear
Handling
to have any lasting effect on the behavior
of foragers. Most
returned within a few
minutes
after their release and went on to make
of
regular trips for the duration
the trial.
on half-hourly
of variance
intervals
Analysis
(Fig. 3) showed no effect of time
=
=
or
treatment
in which
the
{P
0.122)
{P
0.120)
(split-plot ANO VA model
2x2
treatment =
whole plot consisted
of 3 replicated
Latin squares, subplot
intervals were sparse, we reanalyzed
the results
time). Because data for half-hourly
for the entire 2 hours of each trial (Latin square design with
using data summed
3 replicates, with location and trial used as blocking
factors; Table
1). There was
no effect of treatment
or location
the effect of treatment was
(Table 2). While
close to significant,
control dishes had more arrivals than experimental,
the op
if recruitment
occurred. There was a significant
posite of what would be expected
trial effect: control dishes yielded more
at location
1 in two
than experimental
out of three replicates.
in the directions
indicated by the bearings of foragers departing
By searching
from the food dishes, we were able to discover
the nests shown in Fig. 1.At most
of our dishes, foragers arrived from at least two of these colonies. Departures
to
nests #109 and #149 could not be distinguished
because they lay in approximately
area. Foragers departing
the same direction
from the experimental
from the baits
could be tracked visually
cases departing
for up to 20 m. In most
foragers flew
m.
and
at
one occasion
nest
without
the
2-4
toward
of
On
directly
stopping
heights
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
Pi?i
Control
Experiment
o
Time
Fig.
dishes
3.
Numbers
during
each
of new
half
of Agelaia
multipicta
foragers
error.
hour. Error bars = standard
(h)
arriving
on
the experimental
and
control
a departing
from the
forager landed for a few seconds on a leaf several meters
to adjust her load. There was no evidence
of any behavior
sug
bait, apparently
was
no
that
evidence
there
the
leaf.
that
she
scent-marked
Likewise,
gesting
on
to
to
scent-mark.
the
baits
landed
nearby
vegetation
foragers returning
11.2 minutes
after the baits were placed
The first forager arrived on average
n = 12). As the numbers
the
out (SD =?10.6;
of individual
foragers exploiting
found themselves
that two or more
the likelihood
dish increased,
experimental
In some cases, when two foragers encountered
together on the bait also increased.
the other
one another on the bait they displayed
with one chasing
aggression,
each other
from that part of the meat. On other occasions
foragers encountered
Because
such aggression.
without
foragers arrived from two or more
displaying
these attacks may have been on non-nestmates.
colonies,
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
VOLUME 68, NUMBER
285
3
1. Results
of baiting
with Agelaia
Table
Each row represents
experiments
multipicta
(Venezuela).
a single bait dish. Replication:
1 set of 2 paired control
and experimental
baits, set out on successive
=
on
1 = control
1 = dish serving as control
dish. Location:
dish; 2
days. Treatment:
experimental
= dish
=
on day 1 (see Fig.
1 = day 1; 2 = day 2. Arrivals
1). Trial:
day 1; 2
serving as experimental
at
two
each
dish
hours.
number
of new foragers
arriving
during
Treatment
TrialReplication
A
A
A
A
B
B
B
B
C
C
C
C
Location
Arrivals
1
2
1
2
1
2
1
2
1
2
1
2
1
1
2
2
1
1
2
2
1
1
2
2
1
2
2
1
1
2
2
1
1
2
2
1
16
10
5
4
22
6
5
8
6
4
7
6
Peru
and marking
did not appear to disturb foragers arriving at the baits.
Capturing
on the control dishes and released at the end of the trial
of those marked
Many
showed up at the experimental
dishes the next day. New arrivals at the experi
mental
returned
often
within
minutes
after their release and most made
dishes
course
the
of
the
trials.
many
trips during
Most of the arrivals came in the early time intervals (Fig. 4), yet the time effect
=
was only marginally
(P 0.051) (split-plot ANO VA, time nested within
significant
were going on, we would
of arrivals to
expect the numbers
day). If recruitment
increase with time. Analysis
of lumped data (Table 3) showed no effect of treatment
or replication
(randomized
complete block design, blocked on pairs of days) (Table
4).
Our baits were exploited
as
of Agelaia
hamiltoni,
by at least four colonies
taken by foragers departing
determined
from the dishes. The
by the bearings
in the vicinity
locations of three of these were known colonies
of our experiment
(Fig. 2). The fourth, which we could not find, lay to the NW of dish location A.
As was the case in Venezuela,
showed aggression when
foragers sometimes
they
encountered
each other on the baits. As in A. multipicta,
there was no evidence
of anything other than direct, non-stop
flights by foragers to and from the baits.
Table
2.
R = 0.983.
Results
of ANO
VA
on
Source
Sum-of-squares
0.930
Replication
Treatment
Location
(rep)
transformed
Mean-square
2
0.465
1
0.945
2.113
3
0.704
1.741
Error0.127
data
DF
0.945
Trial
1.581
Replication*Trial
root
square
1
1.581
2
0.870
2
0.063
7.345
for Venezuela.
F-Ratio
P
0.120
14.934
0.061
11.129
0.084
24.978 0.038
13.749
0.068
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
Squared
multiple
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
P?I
Control
Experiment
3H
CO
2H
Time
Fig.
dishes
4.
Numbers
during
each
of new
of Agelaia
hamiltoni
foragers
= standard
error.
half hour. Error bars
(h)
arriving
on
the experimental
and
control
after the bait was
first A. hamiltoni
forager arrived on average 2.8 minutes
=
=
n
to
A. hamiltoni,
of
similar numbers
?3.7;
exposed
4). In addition
(SD
to the baits. These wasps are smaller than A.
pollens were attracted
Angiopolybia
to chase
and were often chased by the latter but they never attempted
hamiltoni
A. hamiltoni.
The
Discussion
and A.
that Agelaia multipicta
Our results failed to reject the null hypothesis
resources.
to concentrated
In Vene
do not recruit nestmates
hamiltoni
protein
were not designed
to rule out the possibility
our experiments
of
zuela, although
a general alerting at the nest to a food resource, for example via food odor carried
to the nest, the fact that total numbers
and control dishes) of new
(experimental
over
not
time suggests that this was not going on.
increase
foragers arriving did
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
287
VOLUME 68, NUMBER 3
Table
3.
dish.
of baiting
Results
trial at one
dish.
=
Arrivals
with
experiment
Replicates
positioned
of new
total number
as
one
hamiltoni
(Peru). Each row represents
Agelaia
=
1 = control
2. Treatment:
dish; 2
experimental
two and a half hours.
arriving
during
in Fig.
foragers
Replication Treatment Arrivals
1
2
1
2
A
A
B
B
6
5
5
5
There
data or our direct
is therefore no evidence,
either from our numerical
that foragers of either species recruit
observations
of the behavior
of foragers,
nestmates
to carrion. We conclude
that foragers of these species do not inform
about the location of resources,
either by communicating
naive nestmates
infor
at the nest, by laying scent trails, or by leading them in flight to the site.
mation
as
were not designed
to test for local enhancement,
our experiments
Although
et
&
has been documented
for yellowjackets
Reid
Fowler,
al., 1995),
1983;
(Parrish
the fact that the numbers
of foragers at the experimental
dishes did not increase
over time suggests that it is not going on.
The fact that foragers from two or more colonies
fed at the same dish in each
area
not
indicates
that
these
do
maintain
experimental
species
foraging territories;
instead, the foraging ranges of adjacent colonies overlap. Thus, the foraging strat
find
egy utilized by these species appears to be scramble competition.
Foragers
food quickly
in their foraging area and exploit
it individually
but quickly,
rather
or try to defend a foraging area or a resource against non
than recruit nestmates
nestmate
congeners.
of significantly more foragers at some of the control dishes than at
comment.
in Venezuela
deserves
in handling
is
dishes
Difference
experimental
new foragers arriving at both dishes were captured using
not a likely explanation:
or repellent
to release any attractive
sub
forceps and so had equal opportunity
stances onto the baits. A second possibility
is a reverse local enhancement
effect,
such that foragers are less likely to land on a resource at which one or more wasps
are already present. Our observations
of wasps coming
into the baits revealed no
of this. Third,
it is possible
that foragers avoid areas of high forager
evidence
into areas of low density. Trapping
density and move
foragers out of the area of
so that new foragers moved
in
the density
the control dish may have reduced
areas and discovered
the control baits. Further research on the
from neighboring
to determine
if this result
of foraging in this species will be necessary
dynamics
is anything more
than statistical artifact.
Why do these species lack the ability to recruit to rich patches of food such as
The
Table
arrival
4.
Results
of ANOVA
on
square-root-transformed
data
for Peru.
Squared
multiple
0.667.
Source
Sum-of-squares
Treatment
Replication
0.250
0.250
Error 0.250
DF
1
1
Mean-square F-ratio
0.250
0.250
P
1.000
1.000
0.250 1
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
0.500
0.500
R
=
288
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
carrion? One possibility
is that during
their evolutionary
social wasps
history
on
to give natural
lacked the genetic variability
selection
material
appropriate
in view of the fact that the swarm-founding
which to operate. This seems unlikely,
a system of chemical
swarms
communication
for guiding
species have evolved
from the natal nest to a new nest site (Jeanne,
that adapt
1981, 1991), suggesting
able traits are present.
A second possibility
is that there may be a social constraint.
there is
Perhaps
a threshold colony size below which
the costs of recruitment
exceed the benefits.
to enable a colony
Numbers
of recruitable
foragers may be too low, for example,
to build up enough numbers
at a resource to defend
it against competitors.
Like
are so big that relying on individual
recruitment may pay only if colonies
wise,
to bring in enough
require extremely
large areas to be covered
foraging would
food. We have no idea what critical colony sizes would be, except that it appears
to be larger than for ants (Beckers et al., 1989).
to store food of this type in the nest
lack of ability of wasps
Third, the apparent
mean
to
that
had
little
fitness
is
be
may
gain
by the ability to recruit nestmates
to it. Although
the larvae could likely absorb protein food at higher rates than
the colony
is ordinarily
able to collect it, the size of this buffer may not be large
to
make
recruitment
Some wasp species collect and store nectar
adaptive.
enough
in the nest, sometimes
in large amounts
is
(Jeanne,
1991). If lack of storability
the constraint,
such species recruit to car
it would be worth knowing whether
sources.
bohydrate
factors almost certainly have played a role. Perhaps
carrion
Finally,
ecological
occurs too rarely for this kind of resource to have been a significant
selective force
on foraging behavior
in epiponine wasps. Alternatively,
the benefit/cost
ratio of
to these resources may be too low for wasps,
recruiting
given the nature of com
to nestmates,
that must
exist for them. The costs of communicating
petition
as the time and energy it would
for example,
take to lay a chemical
measured,
can result in effective defense
of the
trail, may not be repaid unless recruitment
or
resource against competitors
same
the
other species. Many
of
species of ants
In fact, some of the ant
to counter.
recruit to carrion and are difficult for wasps
are
at
to
encounter
well
carrion
pose a threat of pr?dation
may
species wasps
likely
for the wasps. Several species of stingless bee are carrion feeders and recruit quickly
the wasps' best strategy may be
(Roubik,
1989). In the face of such competition,
to discover a newly-available
to be opportunistic
i.e.
first
solitary foragers,
by being
it rapidly until ants and/or bees recruit enough
food patch and by exploiting
to force the wasps out.
numbers
on
of foraging in polistine wasps will require proceeding
A full understanding
two fronts. First, we need to extend behavioral
studies such as this one to other
or
to protein
absence
of recruitment
the
to
determine
and
presence
genera
species
studies to learn how wasps
and carbohydrate
food. Second, we need ecological
in
in a variety of habitats
for resources
competitors,
compete
against potential
as
ants
bees
well.
and
not
but
other
wasp
species,
stingless
cluding
only
Acknowledgments
Our thanks go to the Fundaci?n
Branger and to the staff of Hato Pinero and
our visits possible. We
for all their help in making
to Expediciones
Amaz?nicas
James
also thank Dr. Klaus Jaffe for his help in arranging the visit to Venezuela.
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
289
3
VOLUME 68, NUMBER
our specimens. We
thank Erik Nordheim
for sta
Carpenter
kindly determined
on
manu
comments
the
and
drafts
of
For
tistical advice.
helpful
early
thoughtful
we
R.
Stanton
G.
and
thank
Bourne,
Braude,
Zuleyma
Tang
script
Godfrey
for Research
Foundation
Research
Martinez.
supported
by the South African
to RLJ,
the
Science Foundation
National
grant #IBN-9222108
Development,
WI
of
and
Life
of
Madison,
Wisconsin,
Sciences, University
College
Agricultural
and the Department
of Biology, University
53706, and by the Office of Research
of Missouri-St.
Louis, St. Louis, MO 63121.
Literature
Araujo,
Cited
de "Gymnopolybia
para o conhecimento
Contribui?ao
S?o Paulo
11:11-16.
do Instituto
Biol?gico
R. L.
1940.
(Hym.).
Arquivos
and J. M. Paste?is.
1989.
J. L. Deneubourg,
R., S. Goss,
ant foraging
96:239-256.
strategy. Psyche
on the behavioral
1978.
A.
of polygynous
Studies
Forsyth,
ecology
Harvard
Massachusetts.
Dissertation,
University,
Cambridge,
Beckers,
v.
Frisch,
K.
The Dance
Language
Gould,
bridge, Massachusetts.
and W.
J. L., F. C. Dyer,
In B. H?lldobler
and M.
1967.
F. Townes.
Lindauer
141-162.
pp.
B.,
H?lldobler,
chusetts.
1985.
(eds.),
York.
New
Fischer-Verlag,
and E. O. Wilson.
and Orientation
1990.
The
Recent
Experimental
Ants.
meridionalis
Colony
size,
social wasps.
226 pp.
of Bees.
Harvard
(Iher.,
1904)"
and
communication
Ph.D.
Unpublished
Cam
Press,
University
in the study of the dance
progress
language.
and Sociobiology,
Behavioral
Ecology
Harvard
University
Press,
Cambridge,
Massa
swarm emigration
1981. Chemical
communication
in the social wasp Polybia
during
29:102-113.
Behaviour
(Olivier). Animal
1991. The swarm-founding
In K. G. Ross
Polistinae.
and R. W. Matthews
Jeanne, R. L.
(eds.), The
Social Biology
of Wasps,
Comstock
pp. 191-231.
Associates,
Ithaca, New York.
Publishing
L. K.
in Costa
and the structure
of stingless
bee communities
1983.
Johnson,
strategies
Foraging
R. L.
Jeanne,
sericea
Rica.
In P. Jaisson
(ed.), Social
Communication
M.
1961.
Lindauer,
Massachusetts.
Insects
Among
in the Tropics,
Social Bees.
pp. 31-58. Universit?
Harvard
University
Paris.
Paris-Nord,
Press,
Cambridge,
Die gegenseitige
bei den stachellosen
Bienen.
Verst?ndigung
41:405-434.
Physiologie
I. Dietrich,
and W. Keidel.
1974.
bei Wespen
der
Maschwitz,
U., W. Beier,
Futterverst?ndigung
Para vespula. Naturwissenschaften
11:506.
Gattung
and S. Yamane.
1990.
of the Vespine
Berlin.
Matsuura,
M.,
Wasps.
Biology
Springer-Verlag,
S.
1995. Necrophagy
wasps
O'Donnell,
by Neotropical
swarm-founding
(Hymenoptera:
Vespidae,
and W. E. Kerr.
M.,
Lindauer,
Zeitschrift
f?r Vergleichende
Parrish,
Reid,
27:133-136.
Epiponini).
Biotropica
M. D., & H. G. Fowler,
1983.
in two sympatric
related behaviours
Contrasting
foraging
and V. germ?nica.
8:185-190.
wasps
(Vespula maculifrons
Ecological
Entomology
B. L., J. F. MacDonald,
& D. R. Ross,
1995.
and spatial dispersion
in protein
Foraging
sea venging workers
of Vespula germ?nica
and V. maculifrons
Journal
(Hymenoptera:
Vespidae).
of Insect Behaviour
8:315-330.
Richards,
Roubik,
1958.
O. W.
(Natural
D. W.
1978.
History),
1989.
The
Social Wasps
of the Americas
Excluding
the Vespinae.
British
Cambridge
University
Museum
London.
Ecology
and Natural
History
of Tropical
Bees.
Press,
Cambridge.
G.
Sarmiento,
savannas
1983.
The
In F. Bourliere
of tropical America.
(ed.), Ecosystems
Else vier, Amsterdam.
pp. 245-288.
World,
5 Edition.
1992.
Version
Illinois.
Statistics,
Systat for Windows:
Inc., Evanston,
Systat,
1978. Foraging
behavior
of ants: theoretical
considerations.
Journal of Theoretical
Taylor, F.
71:541-565.
Wilson,
E. O.
1971.
The
Insect
Societies.
Harvard
University
Press,
Cambridge,
This content downloaded from 152.1.172.199 on Tue, 21 May 2013 11:12:21 AM
All use subject to JSTOR Terms and Conditions
of
the
Biology
Massachusetts.