New Zealand Journal of Zoology

ISSN: 0301-4223 (Print) 1175-8821 (Online) Journal homepage: https://www.tandfonline.com/loi/tnzz20

Effect of introduced social wasps on the standing

crop of honeydew in New Zealand beech forests

H. Moller, J. A. V. Tilley, B. W. Thomas & P. D. Gaze

To cite this article: H. Moller, J. A. V. Tilley, B. W. Thomas & P. D. Gaze (1991) Effect of
introduced social wasps on the standing crop of honeydew in New Zealand beech forests, New
Zealand Journal of Zoology, 18:2, 171-179, DOI: 10.1080/03014223.1991.10757964

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New Zealand Journal of Zoology, 1991, Vol. 18: 171-179
0301-4223/91/1802-0171$2.50/0 ©Crown copyright 1991
Effect of introduced social wasps on the standing crop of honeydew
in New Zealand beech forests
H.MOLLER 1
J. A. V. TILLEY
B.W.THOMAS
P.D.GAZE2
DSIR Land Resources
Private Bag, Nelson, New Zealand
1Present address:
Zoology Department, University of
Otago, P.O. Box 56, Dunedin, New Zealand.
2Present address: Department of Conservation, Private
Bag, Nelson, New Zealand.
Abstract Honeydew drops produced by beech scale
insects (Ultracoelostoma assimile) on beech trees
(Nothofagus) were measured on three areas of trunk:
an unprotected control area, an area screened to allow
insects but not birds to feed on the honeydew, and an
area screened to exclude all insects and birds. The
experiments showed honeyeater birds (tui, bellbirds,
and silvereyes) had little effect on the honeydew, but
that cropping by German wasps (Vespula germanica),
and particularly by common wasps (V. vulgaris), in the
summer and autumn months markedly reduced the
number, size, and sugar concentration of the honeydew
drops. The energy in drops per unit area of tree trunk
was reduced by at least 99.1 %, 98.4%, and 91.3% in
January, March, and May, respectively. Wasps
reduced the standing crop (number, size, and sugar
concentration) of the honeydew drops by cropping
them before they could be fully recharged and before
evaporation could concentrate the sugars. Cropping
by feral honey bees (Apis mellifera) may have reduced
the standing crop of honeydew slightly. Removal of
the honeydew by the introduced social wasps threatens
the existence of some New Zealand native animals.
The wasps reduce honeydew abundance and quality
to the extent that honey bees do not feed on it for 3-4
months of the year.
Z91011
Received 15 April1991; accepted 17 April1991
171
Keywords Honeydew; wasps; beech; standing crop;
conservation; competition
INTRODUCTION
Honeydew produced by the beech scale insect
(Ultracoelostoma assimile) (Homoptera:
Margarodidae) is an important food of native birds
and insects living in beech forests (Nothofagus spp.)
of the South Island, New Zealand (Gaze & Clout
1983; Clout & Gaze 1984; Moller & Tilley 1986;
Boyd 1987; Beggs & Wilson 1987, 1991; Beggs
1988). In the late summer and autumn of some years,
thousands of introduced German wasps (Vespula
germanica) and common wasps (V. vulgaris)
(Hymenoptera: Vespidae) crawl or hover over the
trunk and branches of each honeydew-covered beech
tree while harvesting the drops or lapping up sugar
spilled on the bark (Moller & Tilley 1986, 1989;
Boyd 1987).
Honeydew drops are smaller and less common
during late January, February, March, and April when
the wasps are abundant (Moller & Tilley 1989). Trees
with more wasps per drop also have smaller drops.
Drops were smaller during summers and autumns in
years of higher wasp abundance (Moller et al. 1988).
These correlations led Moller & Tilley (1989) to
hypothesise that wasps were the main cause of the
reduction in number, size, and perhaps sugar
concentration of the honeydew drops in late summer
and autumn. This hypothesis was supported by a
small-scale exclosure experiment by Beggs & Wilson
(1991). The present study sought to test this hypothesis
by excluding wasps from sections of tree trunks and
measuring the consequent changes in honeydew
abundance and quality throughout the year on several
trees.
Honeydew is the main energy source from which
honey bees (Apis mellifera) (Hymenoptera: Apidae)
make honey in beech forests. Feral colonies now
often inhabit hollow trees in the honeydew beech
forest. and managed hives are kept in some areas to
harvest honey made from honeydew (Anon. 1979;
172
Crozier 1981; Moller & Tilley 1987). The potential
impact of honey bees on the honeydew resource, and
thereby on the native birds that use it, has received
little study. The exclosure experiments reported here
investigated whether feral honey bees affected the
honeydew. The wasps may compete with the honey
bees for the drops, thereby affecting the economic
return from beekeeping in the beech forests
(Clapperton et al. 1989).
Native birds feeding on honeydew are bellbirds
(Anthornis me/anura), tui (Prosthemadera
novaeseelandiae), and silvereyes (Zmterops latera/is).
The main thrust of the present study was to test
whether wasps reduce the standing crop of honeydew
available for the birds to feed on. However, we also
sought to estimate any effect that the birds themselves
might have on honeydew drops, by excluding them
from areas of trunks while allowing the wasps and
bees access.
STUDY AREA
The Trass study area is a 7.2 ha remnant of beech
forest in a pine (Pinus radiata) plantation, 35 km
south-west of Nelson. Red beech (Nothofagusfusca),
hard beech (N. truncata), silver beech (N. menziesil),
and black beech (N. solandri var. solandn) are all
present, but black beech predominates. All except
silver beech have honeydew. The overall abundance
of honeydew at Trass is typical of that in the Nelson
region (Moller et al. 1988). A more detailed
description of the study area. its vegetation, and its
birds is given by Moller & Tilley (1989). There are
no managed hives of honey bees at Trass but at least
four feral colonies occupied cavities in tree trunks in
the area during the study. Nearly all the wasps present
were common wasps (Sandlant & Moller 1989).
METHODS
Exclosures
Ten black beech trees which were heavily infested
with scale insects were used for exclosure
experiments. The lower 3 m of each tree had (i) a
"control'' area giving free access to all wasps, honey
bees, and birds, (ii) a "partial exclosure" area from
which birds were excluded, but wasps and honey
bees could gain access, and (iii) a "complete
exclosure" area, where birds and the insects had no
access to the honeydew. The control area was always
positioned at the top because birds may not have
been as likely to feed underneath the cages used in
New Zealand Journal of Zoology, 1991, Vol. 18
the exclosures. The relative position of the partial and
complete exclosures was alternated between trees.
Exclosure cages were each made from two wire-
mesh half-cages which clipped onto the tree trunk
and then swung shut to join around the trunk. Each
half-cage was 600 mm high, 300 mm wide, and 120
mm deep. The wire mesh was 4 mm square on the
complete exclosure, small enough to exclude wasps,
honey bees, bumble bees, and flies. The partial
exclosure had the same mesh on the front but a 30
mm-diameter round mesh covering the sides; this
was large enough to let insects in but small enough to
exclude tui and bellbirds. Silvereyes could potentially
pass through this mesh, but were never seen to do so.
The exclosures closed tightly against strips of foam
rubber 50 X 50 mm thick inserted between the tree
trunk and the wire frame.
In the first experiment (November 1986) the wire
cages for the complete exclosure successfully
excluded insects. In Jan 1987, wasps found their way
into the exclosure through minute gaps between the
foam rubber strips and irregularities in the OOrk surface.
Accordingly, the January experiment was restarted
with an additional 1 mm square mesh cloth stretched
over the wire frame. This additional barrier was drawn
tightly against the foam rubber strips with a draw-
string, and was used in all subsequent experiments.
Honeydew measurements
The control area and the exclosures each had five
contiguous 100 mm X 400 mm quadrats marked
permanently within them. Each quadrat ran around
the tree trunk, rather than lengthwise, so as to equalise
any effect of aspect on the honeydew between quadrats
(Crozier 1978). On each visit to the tree the drops
within a designated quadrat were counted, and some
of them were collected in a capillary tube to measure
the average drop volume (1 mm column length within
the capillary tube= 1 J..Ll of honeydew). A hand-held
refractometer was used to measure sugar concentration
in grams per 100 g of solution (the Brix scale; see
Bolten et al. 1979). These refractometer readings
reliably estimated the sugar concentration of the
honeydew measured by chemical analysis of its separ-
ate constituents (Grant & Beggs 1989). Sometimes
the drops were too few or too small for sufficient
honeydew to be harvested from the quadrat to measure
the refractive index or even the drop volume.
Sampling regime
We operated the exclosures each alternate month,
beginning in November 1986, and ending in July
1987. On the first day of each experiment we measured
Moller et al.-Effect of wasps on honeydew crop 173

the honeydew in one quadrat in each of control,
partial exclosure, and complete exclosure areas for
all10 trees. We then immediately closed the cages to
exclude insects and birds. We returned 3-5 days later
(Table 1) and measured the honeydew on each tree
from a second quadrat in each area. Cages were only
opened briefly to collect honeydew for measurement
After 2 or 3 days we repeated this procedure, except
that we measured honeydew from a different (third)
quadrat. Again, 2-4 days later, we measured
honeydew on another (fourth) quadrat within each
area and then removed the cages altogether from the
partial and complete exclosure areas. We returned 3-
10 days later to measure the honeydew on the fifth
and final quadrat within each area. This sampling
regime allowed comparison of honeydew in
previously uncropped areas before, during, and after
the experimental treatment
When we visited each tree to measure honeydew,
we counted the number of honey bees and wasps
crawling or hovering over the block of five contiguous
quadrats on the control and partial exclosure area,
and the number of any wasps or honey bees that had
entered the complete exclosure.
We tried to pick dry periods for the experiments,
but sometimes unforecast rain fell or the weather
changed towards the end of the 2-week experimental
time. Rainfall was measured using a Lambrecht
automatic rain gauge just outside the forest on the
study area.

Table 1 Sampling regime and comparisons of honeydew on the control, partial exclosure, and
complete exclosure areas during experiments. Kruskal Wallis tests evaluated the null hypothesis that
honeydew was the same on each area on the given day. In all instances of significant difference the
number, size, and sugar concentration of drops was higher on the complete exclosure area than on the
control and partial exclosure area.
Kruskal Wallis tests
Exper. Rain- No. Drop Sugar
No. Date Sequence fall+ drops size concentration
14 Nov 86 Before ns ns ns
19Nov During 2.1 * ns ns
21 Nov During 0 * * **
25Nov During 7.1 ns ns ns
2Dec After 15.2 ns ns ns
2 16 Jan 87 Before ns ns
19 Jan During 2.1
21 Jan During 0 **
23Jan During 27.8 *** **
27 Jan After 11.8 ns
3 6 Mar 87 Before ns
9Mar During 0 ***
12Mar During# 76.9 ***
4 8 May 87 Before ns ns ns
11 May During 0 * *** ***
13May During 0 ns *** **
15May During 0 ns *** *
25 May After 117.0 ns ns
5 28 Jul87 Before ns ns ns
31 Jul During 0 ns ns ns
2Aug During 8.8 ns ns ns
4Aug During 1.8 ns ns ns
7 Aug After 40.8 ns ns ns
ns P >0.05; * P <0.05; ** P <0.01; *** P <0.005.
#Abandoned because wasps got into the exclosures, it rained heavily, and there was insufficient
honeydew on the control and partial exclosure areas to be measurable.
+ mm rain since previous visit to exclosures.
-untestable because of insufficient honeydew for drop size or refractometer measurement, i.e., there
were fewer than five measurements in the complete exclosure, or in the partial exclosure and control
areas together.
174 New Zealand Journal of Zoology, 1991, Vol. 18

50

40
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160

(/)
c. 100
0

"'E 30 -o
Q; 0 80
~
c. Q)
.0
g 20 E 60
•

~
(/) ::>
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~"' 40
10

0 dJ 20

0
.h ....
Control Partial Complete
ex closure exclosure 0 5 10 15 20 25 30
Number of wasps
Fig. I Mean (±95% confidence interval) number of wasps
per m 2 within treattnents during 35 visits to the experimental Fig. 2 Number of honeydew drops per 100 mm X 400 mm
trees in January 1987. quadrat versus the number of wasps accidentally trapped
within the complete exclosure area in March 1987.

Non-parametric statistics have been used to test

null hypotheses in most instances, because many of
the insect counts and honeydew measurements are
distributed non-normally.
RESULTS
Did the exclosures work?
No birds were ever seen feeding on the quadrat areas
when the cages were shut. Bellbirds were seen feeding
on honeydew very close to the side of the exclosure
cages, so we have no reason to suspect that the
control area was avoided by the birds because of its
proximity to the exclosure structures.
Wasps and honey bees did feed behind the wire
cages on the partial exclosure, but not to the same
extent as on the control area. For example, in January
1987 there were only about a third as many wasps
feeding on the partial exclosure area as on the control
area (Fig. 1). Wasps and honey bees were often seen
to fly up to the open mesh cages, hover 10-20 mm
from the wire, and then veer off. Most of the honey
bees and wasps feeding on the partial exclosure area
had crawled or flown through the large gap on the
side of the cage.
Despite the outer mesh cloth added to the complete
exclosure from January onwards, wasps were
sometimes able to crawl through minute gaps in the
seal (Fig. 1). The honeydew was much reduced in
exclosures that wasps had forced their way into (Fig.
2). Measurements from these exclosures were
eliminated from the analyses which follow.
Insect and honeydew measurements
on the control area
Feeding wasps were abundant on the honeydew-
producing trees in January, March, and May but
virtually absent in November and July (Fig. 3). One
tree in March had the equivalent of 370 wasps per m2
of trunk crawling over the control area; the average
for that month was 119.8 (n = 30, SE = 14.4) wasps
per m2. In contrast, feral honer bees never averaged
more than about one bee perm of bark (Fig. 3). None
were observed on the trees in March when the wasps
were at peak abundance.
So little honeydew was present in January and
March that refractive index and drop size could not
be measured. There were too few drops, and they
were too small in January and March to form a
column in our capillary tube, even though average
drop sizes of about 0.1 J1.l. are detectable by the
technique (e.g., in May, Fig. 5). Hence, the average
drop size in January and March on the control area
and the partial exclosure was less than 0.1 Jl.l..
Number and size of drops declined between
November and January as the wasps appeared, were
lowest in January and March (Fig. 4 and 5) when
wasps were highest (Fig. 3), and then increased again
by May when the wasps were nearly gone.
Honeydew within the exclosures
There were no significant differences in the number
of honeydew drops, their median size, or their sugar
Moller et al.-Effect of wasps on honeydew crop
160
120
I
,j,
' • Wasps
I '
/ \
'
I
''
' similar to, or slightly higher than on the control area.
I '
40
0 0
NOV JAN MAR
1966 1967
Fig. 3 Mean (±95% confidence interval) wasp and honey
bee density on the control area during each experiment.
concentration on the quadrat areas before any of the
exclosure cages were attached (fable 1). Similarly,
any differences in honeydew which developed during
the period of exclosure disappeared again after the
cages were removed.
Whenever there were significant differences
between treatments, the drop number, size, or sugar
concentration were always lower in the control and
partial exclosure than in the complete exclosure area.
Differences in the size of drops were greatest in
January and March, when wasp abundance was
greatest. Sugar concentration was significantly lower
within the control and partial exclosure than in the
complete exclosure in November and in May (fable
1, Fig. 6), when a comparatively small number of
wasps and honey bees were harvesting the honeydew
drops. This suggests that sugar concentration would
also have been markedly lower in the control and
partial exclosure area than in the complete exclosure
area in January and March, when insufficient
honeydew could be collected to measure it. The
decline in sugar concentration between May and JulyI
August on the control area (Fig. 6) was probably
caused by rain, which fell just before our 2 August
sampling (fable 1).
We calculated the standing crop of honeydew
(energy value per unit area of tree trunk) in the
complete exclosure and on the control area for the
middle day of each exclosure experiment by (1)
assuming that honeydew in January and March would
have had a concentration of 45 g per 100 g, had we
175
been able to collect sufficient of it to measure, and (2)
assuming that the average drop size in January and
March was 0.05 Jll. The standing crop on the control
area was0.8%,1.6%,and9.7% of that on the complete
exclosure in January, March, and May, respectively,
o Bees
the months when wasps were abundant.
The number, size, and sugar concentration of
1·6 honeydew drops on the partial exclosure tended to be
1·2
g>
~
o-6 " DISCUSSION
~
0·4 Effects of exclosures
Honeydew on the control and exclosure areas was
always about the same before and after the
MAY JUL experimental treatment. This suggests that the relative
position of the quadrats on the tree had little impact
on our results. Beggs (1988) found that honeydew
drops became more concentrated on the branches of
the tree than on the lower trunk.
The appearance of significant differences in
honeydew once the exclosure cages were closed, and
their disappearance after removal of the cages, leaves
no doubt that the treatments caused the observed
changes in the honeydew. However, the changes in
honeydew might have resulted in part from changes
in microclimate and the impact of weather on the
enclosed honeydew, rather than simply from the
elimination of cropping of the drops by the insects
and birds. Weather effects might include:
(a) the thick foam-rubber strips sealing the gap
between the exclosure cage and the bark probably
reduced the amount of rainwater diluting the
honeydew by streaming down the tree trunk and
down the anal tubes of the scale insects within the
exclosure area;
(b) reduction of wind and sunlight within the
exclosures might have increased the relative
humidity within the exclosure, and thereby
decreased the rate of evaporation of water from
honeydew drops;
(c) screening by the cages might have reduced the
rate at which drops (of a given size and sugar
concentration) are knocked or blown off the anal
tubes.
Factor (a) would increase sugar concentration
and reduce drop size in complete exclosures. Factor
(b) would tend to cause sugar concentration and drop
size to be reduced within the complete exclosure, but
this effect might be partially or completely
counteracted by (c) because less knockdown of drops
would lead to them remaining on the anal tubes for
176
70
60
9
~50
"'cr
::>
:;; 40
"-
"'
"-
~ 30
0
20
10
longer (drop size and sugar concentration increase
with the time since the drop first began to fonn-see
later). Beggs & Wilson (1991) found that honeydew
within exclosures (similar to those used here) remained
more concentrated than on control areas even after
the wasp numbers declined. However, Beggs &
Wilson left their exclosures pennanently shut for
months. In our shorter experiments, we found no
differences in honeydew when wasps or honey bees
were in low numbers (Table I, Fig. 4--6).
There are three other indications that changes in
honeydew in our experiments were brought about
more by eliminating cropping than by microclimate
or weather-screening effects:
(1) Differences developed in honeydew between
treatments even in dry weather.
(2) During the peak wasp season, large honeydew
drops appeared within 20 min of the closing of
the complete exclosure. This response was so
rapid that it must have been related to cessation of
cropping by wasps and not the influence of
microclimate within the complete exclosure.
(3) Honeydew was much reduced within the complete
exclosures whenever wasps found their way in
through minute gaps in the seal or were
inadvertently trapped inside when the cage was
closed (Fig. 2). The gaps were so tiny that they
could not have altered the microclimate to any
great extent, so the reduction of honeydew in
New Zealand Journal of Zoology, 1991, Vol. 18
9Control Fig. 4 Mean (±95% confidence
interval) number of honeydew
+Partial drops per 100 mm X 400 mm
9Complete quadrat in each treatment area on
the ftfth or sixth day after closing
the exclosure cages. The sample
size is given by each point when
honeydew was measured on less
than 10 trees.
such accidental "enclosures" must have been
caused by trapped wasps cropping the drops.
Effects of birds on honeydew
The reduction in the number of wasps and honey
bees feeding on the partial exclosure area compared
to the control area (Fig. I) means that our experiment
failed to separate completely the effects of birds and
insects on the honeydew. Despite this, the amount
and quality of the honeydew on the partial exclosure
area was similar to that on the control area (Fig. 4--6).
This suggests that (i) the insects were so abundant
even on the partial exclosure area that the increase in
their density on the control area made negligible
impact on the relative amount of measurable
honeydew there, and (ii) that feeding by birds had
relatively little effect on honeydew compared to
feeding by insects.
Effects of wasps on honeydew
The exclosures would have prevented bumble bees,
large flies, and some ants from feeding on the
honeydew, as well as wasps and feral honey bees.
However, in the summer and autumn of 1986 and
1987 wasps far outnumbered the other insects, and
honey bees predominated in the spring and winter
(Moller & Tilley 1989). Wasps even fed on the
honeydew throughout the night (unpubl. obs.). In
1988 when the abundance of wasps was an order of
Moller et al.-Effect of wasps on honeydew crop
Fig. 5 Mean (±95% confidence 09
interval) size of honeydew drops in
each treatment area on the fifth or
0·8
sixth day after closing the ex closure
cages. The sample size is given by
each point when honeydew was
measured on less than 10 trees.
magnitude less than in the two previous years, they
were still the most numerous honeydew feeders
(Moller et al. 1988). This suggests that during our
experiments, and in most years, wasps have a major
influence on the number and quality of honeydew
drops.
Wasps cropped drops so frequently at the peak of
the wasp season that we could not collect sufficient
honeydew to compare the size and sugar concentration
of drops inside and outside the exclosures. However,
the increase in drop size in the complete exclosure in
January and May and in sugar concentration in May
leaves no doubt that wasps reduce not only the number
of drops available for other feeders, but also the
average sugar content.
Comparisons of honeydew on the control and
complete exclosure areas suggest that the wasps had
reduced the standing crop of energy (in drops) by
99.1 %, 98.4%, and 91.3% in January, March, and
May, respectively. These comparisons are likely to
under-estimate the impact of wasps on the resource
because (i) the standing crop of honeydew tends to
increase in the absence of rain and cropping by the
wasps, so the comparison in standing crop between
treatments for later in each experiment would also
have revealed greater differences between treatments,
and (ii) we used safe, but high, estimates of drop size
and sugar concentration in January and March, so as
to avoid overemphasising the effects of the wasps on
the honeydew resource.
177
9Control
+Partial
9Complete
Effects of honey bees on honeydew
There were significant but slight differences in
honeydew between treatments in November (Table
1). However, even a slight interference to
microclimate within the exclosures, or screening
from rain (see above) may have caused the slight
differences between honeydew in the different
treatments in November. Accordingly, we cannot
conclude that the differences definitely resulted
from the effects of cropping by the feral honey bees
living on our study area. Clearly any impact of feral
honey bees on the honeydew is much less
pronounced than that of the wasps.
Drop recharging-the mechanism by which
wasps affect honeydew
When a drop is removed from an anal tube a new
drop of substantial size is sometimes formed within
minutes. Some tubes take much longer to form a new
drop, and this is reflected in the gradual increase in
drop numbers within the complete exclosure as each
experiment progressed. Moller & Tilley (1989)
showed that mechanically stimulating the anal tube
or tapping the scale insect's capsule sometimes
triggered the rapid formation of a drop on an anal
tube. As more fluid accumulates at the tip of the anal
tube the drop grows in size. Evaporation of water
from the drops eventually concentrates the sugar to
the extent that the drops resemble toffee.
178 New Zealand Journal of Zoology, 1991, Vol. 18

9Control Fig. 6 Mean (±95% confidence

interval) sugar concentration of
+Partial honeydew drops in each treatment
area on the fifth or sixth day after
6 ?Complete closing the exclosure cages. The
'2
,g 60 sample size is given by each point
0" when honeydew was measured on
"'
g55
less than' 10 trees.
0 9
....
3
.§ 50
~
"~
c
0
45
0

~ 40
"
en

NOV JAN MAR I I MAY I JULIAUGI

1986 1987

Our exclosure experiments show that wasps (and
perhaps also honey bees to a small extent) can lower the
number, the size, and sugar coocentration of drops by
revisiting them ~ often that drops do not last long
enough to I'€'1JCh maximum size and sugar coocentration.
Competition between wasps, honey bees,
and birds
Feral honey bees ceased feeding on the honeydew
during the peak of the wasp season, probably because
feeding on the few small remaining drops was un-
profitable. The year after the present study there were
com{m3tively low numbers of wasps, more honeydew
and more feral honey bees feeding on the trees
throughout the late summer and autumn (Moller et al.
1988). This suggests that there is feeding competition
between the honey bees and the wasps. The removal
of from 91% to over 99% of the honeydew standing
crop by the wasps, as demonstrated by the exclosure
experiments, suggests that competition for this main
food is the cause of the interaction. It also corroborates
several indications that honeydew is a limiting
resource for wasp populations (Moller et al. 1990).
There are no other abundant sources of nectar or
honeydew in the study area (and in most areas of
beech forest) to provide alternative foods for the
wasps or honey bees. Competition for this limited
resource is probably also important in the complete
competitive exclusion of German wasps upon the
recent arrival of common wasps in these honeydew
beech forests (Moller et al. 1990; Harris et al. 1991).
Virtual elimination of the honeydew standing crop
for 3-4 months of the year may affect the native birds
that feed on the drops (Gaze & Clout 1983; Clout &
Gaze 1984; Beggs & Wilson 1987; Moller & Tilley
1989; Beggs & Wilson 1991).
ACKNOWLEDGMENTS
Work on the last two exclosure experiments was funded
by the Department of Conservation. We are also grateful
to Baigent Forest Industries Ltd for allowing us to use the
Trass study area; and to the Canterbury Beekeepers
Association and the Honeydew Producers Association for
providing a refractometer for the study. Merle Rae typed
the manuscript, which was improved by criticisms from
Jacqueline Beggs, Tony Pritchard, Chris Thomas, and
particularly by Murray Efford.
REFERENCES
Anon. 1979: Beech honeydew honey-a vast potential.
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