Preliminary Observations on Foraging Activities of Bombus dahlbomii

and Bombus terrestris (hym: Apidae) on Native and Non-Native

Vegetation in Chile
Luisa Ruz1,3 and Raúl Herrera2,3
1
Laboratorio de Zoología
2
Instituto de Estadística
3
Universidad Católica de Valparaíso
Av. Brasil 2950
Valparaíso, Chile.

Keywords: Bumblebees, foraging, pollen, nectar, Eryngium, Lavatera, Salvia

Abstract
The present paper provides preliminary information about some foraging
activities of B. Dahlbomii, a native apid, and the introduced B. terrestris, in areas of
Central-South and Central Chile. The field work was centered in looking at
variables like: frequency of insect visits per plant; stigma contact; pollen collection;
nectar gathering; and, time of floral visitation. Analysis of the data attempts to
provide a better understanding of bumblebees behavior when foraging on native and
non-native flowering plants and interaction with other insects, especially bees, in
wild conditions.

INTRODUCTION
Bombus dahlbomii Guérin. Is a well-known, native bumblebee of impressive size
and beauty and in the past was very common (Montealegre, 1927), frequently found in a
geographical range from Central to Southern Chile. Over the last 25 or 30 years however,
populations of this species have apparently decreased. This statement still needs to be
quantified with the introduction of B. terrestris in 1998 for tomato pollination in
greenhouses (Beèche, pers. Communic.). It is especially necessary to conduct studies
leading to a better knowledge and understanding of these species, their behaviour, and
their interactions with flowering plants and other bees under wild conditions.
The need for more information about the present situation of B. dahlbomii and of
the possible factors influencing its population size motivated us to conduct this study.
Also under consideration was B. terrestris, one of the most aggressive bumblebees known
and concerned about endemic pollinator displacement (Stephen, pers.com.). This latter
species has proven to expand rapidly and act negatively with other bees in Israel (Dafni
and Shmida, 1996). In Chile, however, no studies have been carried out on B. terrestris in
the wild, except those involving its rearing under laboratory conditions (Wagner,
pers.com.) and its role as tomato pollinator in greenhouses (López, pers.com.). On the
other hand, even though the presence of B. terrestris has had positive effects on
bumblebee research and has been successfully ommercialised, its introduction to new
areas is risky and the consequences for the environment unknown (Ruijter, 1996).
Because of the polylectic behaviour of bumblebees, it would seem imminent that
in Chile B.dahlbomii and B. terrestris, during overlapping periods of their life cycle,
would visit the same floral resources, with the latter likely outcompeting the first or other
native solitary bees. The main goal of this preliminary study is gathering behavioural data
related to foraging activities of B.dahlbomii and B. terrestris, as well as to some
interactions with flowering plants and other bee species under wild conditions.

MATERIALS AND METHODS

The observations were made in the following localities from two main Regions: a)
Cobquecura, long.36º 07’ 38S, lat.72º 48’ 14W, in Central-South Chile (VIII Region); b)
Curauma (Valparaíso), long.71º 39’ W, lat.33º 04’ S and Lilenes-Con-Cón (Viña del
Mar), long.71º 31’ W, lat.32º 56’ S; and c) Putaendo, long.70º 43’ N, lat.32º 38’ S at

Proc. 8 th Pollination Symp.

Eds. P. Benedek & K.W. Richards
Acta Hort. 561, ISHS 2001 165
almost 1000 m altitude (V Region). The field work was carried out during the periods:
January 18 -25; April 26-27, 30; and May 1,7, 2000.
At the study sites behavioural observations and censuses of insects visiting the
plants were done mostly by sight and collecting with entomological net was minimum.
In order to achieve the main objective of this study and to know specific
behaviours, the following variables were recorded in the field and then statistically
analyzed: species of insect; daily activity hours; time of floral visitation; stigma contact;
nectar collecting; pollen collecting; species of plants; and, abiotic factors, such as
temperature and relative humidity. These data were analyzed by one of the authors (R.H.)
using the English Version 8.0 for Windows of the software SPSS. The censuses were
done daily for five minutes, every hour, from 10:00 am to 6:00 pm. During each census,
insect individuals were recorded each time they arrived at a flower. For recording time of
floral visitation, the insects, especially bees, were observed until they flew away. Pollen
loads were observed and grains identified; nectar contents of some flowers were also
measured and analyzed. Insect and plant species were identified in the Laboratory.

RESULTS AND DISCUSSION

Forager Records and Food Resources

Among the Apoidea found in the study sites as a whole (92.1%), B.dahlbomii was
the most frequently observed (freq=812, almost 50%). In Cobquecura, more than 50%
corresponded to this species, of which almost 50% visited the native plant Eryngium
paniculatum (Apiaceae). Lower frequencies of B.dahlbomii were observed on: Raphanus
sativus (Brassicaceae); Cirsium vulgare (Asteraceae); Mentha pulegium (Lamiaceae);
and, Trifolium striatum (Fabaceae). In Curauma, B.dahlbomii was less common, visiting
only Lavatera arborea (Malvaceae) (freq=274) and was absent in Lilenes, even though
Eryngium and Lavatera were present (the last two areas in the coast). B. terrestris
corresponded only to 10% of the total and was found in Putaendo (freq=170) and mostly
observed visiting Salvia leucantha (Lamiaceae), an ornamental introduced species; a few
times it was seen on Geranium sp.(Geraniaceae) and on Chrysanthemum sp2
(Asteraceae); another 3 plant species present in the area were not visited. The numerical
differences between records of both bumblebee species, however, are not comparable.
The observation period for B. dahlbomii was in January, while the colonies were still very
active and flower abundance was high, while B. terrestris was, for the first time, found in
the wild during April-May, when the colony was small and the queens were looking for
suitable hibernation places. The abiotic factors where these species were present differed;
Putaendo had the lowest temperature (10ºC) and highest relative humidity (90%). These
climatic conditions favoured B. terrestris activities. While the other 3 localities together
present a T range of 15.9°-28.6°C and a relative humidity between 34.4% and 56%.
Foraging Activities

1. Stigma Contact. Through observation, several insects were detected, in general terms,
as good pollinators of plants because of the high proportion of contact with the floral
stigma. The most remarkable corresponded to C. bicolor, a native bee, with almost 91%
contacts. In B. dahlbomii, contacts diminished to 35%; this species mainly touched the
stigma of M. pulegium, T. striatum and L. arborea; however, not that of Eryngium, which
flowers were too small compared to the large-sized, long-legged bee. B. terrestris
contacted the stigma of Geranium sp., but not that of Salvia leucantha. This behaviour
depends on the floral and insect features like size and morphology.

2. Nectar Harvested. Both B. dahlbomii and B. terrestris were almost exclusively

collecting nectar (Table 1). B. dahlbomii was observed robbing nectar from Lavatera
arborea (ornamental plant) in Curauma, but not from the rest of the plants from
Cobquecura. B. terrestris, on the other hand, was always observed “robbing” nectar from

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Salvia leucantha, and hence was not an efficient pollinator. Samples of nectar were taken
from 60 flowers at 5:00pm with an average of 1.9 ml/fl. The sugar content of 80 mg was
analyzed and the following approximate values were obtained (mg): glucose 1.98,
fructose 0.25, sucrose 10.83, maltose <0.1. Nectar analysis for Eryngium and Lavatera
failed because of extraction difficulties. Flight patterns were not very consistent and need
to be clarified.

3. Pollen Gathering. Pollen loads were observed on solitary bees (i.e. Callonychium
chilense, Colletes cognatus), but not on females of B. dahlbomii. Except when they
visited Eryngium, were pure and abundant pollen grains found on their body, an important
fact from the pollination viewpoint, although individuals were not observed touching the
stigma; only in one case (not recorded) pollen load was identified as being from
Eryngium. Their medium-sized pollen grains fall into the range 25-50µ established by
Erdtman (1952). This size contrasts strongly with those of Lavatera arborea (98-107µ),
which had conical spines of 8µ length, according to Pla (1961). Little pollen was carried
by B. dahlbomii from Lavatera (heavy loads were never observed) and may be explained
in a similar way to that of honeybees, which present reduced efficiency to aggregate spiny
pollen grains from Gossypium (also Malvacea) (Vaissière and Vinson, 1994); although
the large size hypothesis of Buchmann and Shipman (1990) may also be true. On the
other hand, females of B. terrestris were never observed collecting pollen from Salvia
leucantha. Two queens, however, were carrying pollen loads from this plant species.

4. Time of Floral Visitation. There is a clear variation in the time required by each
species per flower visitation (Table 2). Comparing both bumblebees, it is evident that B.
terrestris is much faster than B. dahlbomii, taking on average only 8.64 seconds per visit
against 20.34 seconds for the latter species. From the information given in the Table, the
number of flowers visited per minute can be estimated for each species. B. dahlbomii, A.
mellifera and C. bicolor can visit near 3 flowers per minute, D. chilensis almost 6, and B.
terrestris around 7. Values of 30 for bumblebee species (doubling those of honeybees)
were obtained by Heinrich (1981). The species with more disperse time data was A.
mellifera and the one with less disperse values (more homogeneous) was B. terrestris
(Figure 1). A Test for Median differences determined a statistically significant difference
between the two species of Bombus with T=9.58 and a p-value close to zero. Other
species were included in the graph for comparisons.

CONCLUDING REMARKS
B. dahlbomii was the only bumblebee detected in Cobquecura (VIII Region), a
study site where the higher percent was found, being the most frequently observed
foraging in Eryngium paniculatum, a native plant also visited by other insect species,
especially bees. Its large size apparently prevented it from touching the stigma, so it
should not be an efficient pollinator of this plant, however it several other plant species.
B. terrestris, detected for the first time in wilderness, foraged faster than
B.dahlbomii and other natives bees, like Colletes bicolor. Therefore, in the long-term, it
may cause an impact over slower forager species when exploiting the same resource.
Since the two bumblebee species were not sharing the same plants nor the same
study sites under equal climatic conditions, further investigation would be needed to gain
knowledge about their behaviour during overlapping activities of their life cycle.

ACKNOWLEDGEMENTS
Many thanks to the following people that kindly collaborated in some way with
this study: Dr. Hugo Monzón, for his suggestions and interesting ideas; the students
Paulina Mena, Felipe Vivallo, Daniel Tapia, Macarena Bugueño and César Badillo, for
gathering data during field work; Mauricio Cisternas, for plant identification. All
collaborators are from the Universidad Católica de Valparaíso, Chile.

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Literature Cited
Buchmann, S. L. and Shipman, C.W. 1990. Pollen harvesting by Apis mellifera on
Gossypium (Malvaceae) flowers. Jour. of Kansas Entomological Society, 63:92-100.
Dafni, A. and Shmida, A. 1996. The possible ecological implications of the invasion of
Bombus terrestris L) (Apidae) at Mt Carmel, Israel. In Matteson, A., Buchmann, S.L.,
O’Toole, C., Westrich, P. and Williams, I.H. (eds.). The Conservation of Bees.
Academic Press, London: 183-200.
Erdtman, G. 1952. Pollen morphology and plant taxonomy: Angiosperms. In “Pollen and
spores of Chile. Modern types of Pteridophyta, Gymnospermae and Angiospermae”
by Heusser,C. J. (1971). The University of Arizona Press. Tucson, Arizona : 1-167
Heinrich, B. 1981. Bumblebee economics. Harvard University Press. Cambridge,
Massachusetts and London: 1- 245.
Montealegre, A. 1927. Biología de insectos chilenos. IV. El Moscardón (Bombus
dahlbomii). Revista Chilena de Historia Natural, 31: 165-172.
Pla, J.M. 1961. Polen. Estructura y características de los granos de polen de especies
recolectadas en el N.E. de España. Polinización y Aeropalinología.Talleres Gráficos
D.C.P. Gerona: 1-508.
Ruijter, A. de 1996. Commercial of Bumblebee rearing and its implications. In Richards,
K.W. (ed), 1997 Proceedings of the 7th International Symposium on Pollination, Acta
Hort. 437,261-269.
Vaissière, B.E. and Vinson, S.B. 1994. Pollen morphology and its effect on pollen
collection by honey bees, Apis mellifera L. (Hymenoptera: Apidae), with special
reference to upland cotton, Gossypium hirsutum L. (Malvaceae). Grana, 33:128-138.

Tables

Table 1. Distribution of records for B.dahlbomii and B. terrestris according to the type of
product collected from plant species

Product collected
Apoidea species Plant species N P N&P Total
E. paniculatum 471 0 2 473
R. sativus 39 0 0 39
C. vulgare 13 0 0 13
Bombus dahlbomii M. pulegium 3 0 0 3
T. striatum 10 0 0 10
L. arborea 119 11 144 274
Total 655 11 146 812
Geranium sp. 10 0 0 10
Chrysanth. sp2 8 0 0 8
Bombus terrestris
S. leucantha 152 0 0 152
Total 170 0 0 170

N = Only nectar; P = Only pollen; N & P = nectar and pollen

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Table 2. Descriptive measures of the floral visiting time used by different insect species.

Insect species N Mean Med. S. D. Variance

B. dahlbomii 812 20,34 12 25,36 643,27
A. mellifera 405 20,22 12 22,54 508,11
C. bicolor 98 18,39 9 26,06 678,90
B. brullei 114 13,07 8 19,26 370,79
D. chilensis 26 14,38 8,5 14,92 222,57
B. terrestris 170 8,64 6 10,93 119,36

Figures

60
Time of visitation on flowers (sec)

50

40

30

20

10

0
N= 812 405 98 114 170

Bombus dahlbomii Colletes bicolor Bombus terrestris

Apis mellifera Bembex brullei

Species of insects

Fig. 1. Time of floral visiting plotted for different species of insects.

169