Journal of Apicultural Research 33(1): 15-20 (1994)
Copulation of Bombus terrestns
L. (Hymenoptera=
Apidae) in
@1994 |BRA
captivity suMMARY
Y D.lecHAM; J C VenrnecxE; P RnsmoNT
Laboratoire de Zoologie, Université de
Mons-Hainaut, 8-7000 Mons, Belgium
(Received 10 March 1993,
accepted subject to revision 29 September 1993, accepted for publication 12 October 1993)
The mating behaviour of Bombus terresfris was studied using wooden observation cages (50 x 50 x 7O cm) in the laboratory. Queens and males from reared colonies were isolated from the odour of the opposite sex before the experiments. Mating pairs (one per box) con- sisted of one 1O-day-old virgin male and one 5-day-old virgin queen. The behaviour of each of the 80 pairs used was observed for 15 min, during which time 47 of the pairs copulated.
Male behaviour could be divided into approach, inspection and attempt to copulate.
Queen reactions to the male were immobility (1 3 .9Vo), threat (53 .9yo) or flight (32.3o/o).
Queens could also display inspection behav- iour. The mating success of a behavioural sequence was positively correlated with the frequency of inspections by both sexes and negatively correlated with queen immobility.
Flow diagrams showing seguences of types of behaviour are presented. The role of sexual pheromones in mating is discussed.
Keywords: bumble bees, Bombus
terrestris, mating behaviour, sexual
behaviour, copulation
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Copulation of Bombus terrestris 1 7
Behavioural frequency analysis
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Approach* Inspect female*** Inspect male*** Attempt to copulate
m Successful [-l Unsuccessful
M - nbr.of act lduration of the sequence (sec.)
FlG. 1. Behavioural frequency analysis. Median frequency of the differcnt behaviours (number of observed/duration of the sequence (s)) for the successful sequences (culminating in copulation) and for the unsuccessful sequences (median test, * = significant, *** = vêr'!f highly significant).
the approach, inspection lasts only few seconds
thus we also consider it as an event.
3. Attempt to copulate (AG): the male grips the
queen, places his middle legs on her abdomen and
tentatively begins periodic genital insertions and
simultaneously touches the queen's thorax with his
antennae. This behaviour lasts from 1 s to 840 s.
Ethogram: female behaviour
As noticed by the authors, the female is far more
passive than the male. When she is not reacting to
an approach of the male, she is usually settled on
t h e cage, immobile.
However, she reacts to the approach of the male in
three ways:
1. lmmobility (O): the queen stays immobile during
and after the male approach.
2. Threat (+): the queen is settled on the cage,
immobile. When the male approaches she raises
towards him her middle legs and moves away. This
behaviour is generally considered as a threat.
3. Flight (-): when the male approaches the queen,
she flies away from him.
Moreoveri the female can also display inspection (lM)
of her partner. She flies towards the male and touch-
es his abdomen and head with her antennae. This
behaviour is identical with that of the male. ln this
behaviour the female flies behind the male with her
antennae pointed towards him.
Frequency analysis
The success of the sequence was correlated with
the frequency of inspections by both sexes (lF and
l M ) (f i s . 1 ) .
Analysis of the female reactions to the approach of
the male (A) (immobility: 13.8%o of her reactions;
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Copulation of Bombus ferresfns 1 9
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cop
FlG. 3. Flow diagram of unsuccessful courtship (the sequence does not end in copulation). The thickness of the arrows is proportional to the probability of the transition from one behaviour to another. The area of the circles is proportional to the frequency of the behaviour. A = approach, lF = inspection of the female by the male, AG = attempt to copulate and COP = copulation. k' test, * = significant, ** = highly significant, *** = vêry highly significant). Total number of observed transitions = 1 029.
attractant and arrestant for the queen. When queens
are inspecting scent sites along the flight routes,
males are thought to be attracted to them and may attempt to copulate. lt seems that the odour of the q u e e n is important in eliciting contact and an attempt to copulate.
As is the case in other insects with low visual acuity,
mistakes made during visual approach can be rec-
tified by scent stimuli at close quarters (Free,1970,
1 9 8 7 ) . T h e odour of the queen comes from a
pheromone produced by the mandibular glands.
In our experiments, unlike the situation in the field,
males did not establish a nuptial route; the space
was too restricted. In other respects, we observed
the same male behaviours as in the field (approach,
inspection and attempt to copulate). Since the male
did not mark scent sites in our experimental cage,
we noticed that he was directly inspected by the
queen. This is the first time this queen behaviour has
been observed.
Previous authors have stressed the importance of
male and female sex pheromones in the courtship
behaviour of bumble bees (Sladen, 1912; Frison,
1927; Frank, 1941; Haas, 1949, 1967; Krûger, 1951 ;
C u m b e r , 1 9 5 4 ; K u l l e n b e r g , 1 9 5 6 ; S t e i n , 1 9 6 3 ;
Awram , 1970; Free, 1970, 1987; Schremmer, 1972,
Bringer, 1973; Honk et al., 1978; Svensson, 1980).
This study conforms with these results since the
antennal inspections seem to be a key factor in the
success of the sequence. However, the analysis of
t h e f l o w diagrams suggests that if the female
p h e r o m o n e e l i c i t s c o p u l a t i o n it d o e s n o t d o s o
immediately: antennal inspection of the female by
the male is never directly followed by an attempt to
copulate. So the sex pheromones, in addition to
their attractive and species-recognition role, seem
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