Aggressive and Docile Colony Defence Patterns in Apis mellifera. A Retreater–Releaser Concept

Abstract

Colony defence in Apis mellifera involves a variety of traits ranging from 'aggressive' (e.g. entrance guarding, recruitment of flying guards) to 'docile' (e.g. retreating into the nest) expression. We tested 11 colonies of three subspecies (capensis, scutellata, carnica) regarding their defensiveness. Each colony was selected as reportedly 'aggressive', 'intermediate' or 'docile' and consisted of about 10,000 bees. We applied three stimulation regimes (mechanical disturbance, exposure to alarm pheromones, and the combination of both) and measured their behaviours by tracking the rates of outflying bees at the entrance sites of the test hives. We provided evidence that for mechanical disturbances the test colonies resolved into two response types, if the 'immediate' defence response, assessed in the first minute of stimulation, was taken as a function of foraging: 'releaser' colonies allocated flying guards, 'retreater' colonies reduced the outside-hive activities. This division was observed irrespective of the subspecies membership and maintained in even roughly changing environmental conditions. However, if pheromone and mechanical stimulation were combined, the variety of colony defensiveness restricted to two further types irrespective of the subspecies membership: six of nine colonies degraded their rate of flying defenders with increasing foraging level, three of the colonies extended their 'aggressiveness' by increasing the defender rate with the foraging level. Such 'super-aggressive' colonies obviously are able to allocate two separate recruitment pools for foragers and flying defenders.

What causes wing wear in foraging bumble bees?

“…Many studies have used wing wear to estimate relative insect age (Mueller and Wolf-Mueller, 1993;Kemp, 2000;Burkhard et al, 2002;Richards, 2003;Inoue and Endo, 2006;Peixoto and Benson, 2008). Wing wear has consequences, which include increased wingbeat frequency (Hargrove, 1975;Kingsolver, 1999;Hedenstrom et al, 2001), changed flight speed (Fischer and Kutsch, 2002), changed flight performance (Haas and Cartar, 2008;Jantzen and Eisner, 2008;Combes et al, 2010), changed foraging behaviour (Higginson and Barnard, 2004;Foster and Cartar, 2011) and increased risk of mortality (Cartar, 1992).Many eusocial insects, particularly bees and wasps, rely on their wings to defend their nest from predators (Breed et al, 1990;Kastberger et al, 2009), maintain colony temperature and assure proper larval development (Heinrich, 1979a;O'Donnell and Foster, 2001), and acquire food for themselves and their colony (Heinrich, 1979a). Providing protection, care and food for the colony's young are the ways in which a non-reproductive forager increases its inclusive fitness.…”

“…Many eusocial insects, particularly bees and wasps, rely on their wings to defend their nest from predators (Breed et al, 1990;Kastberger et al, 2009), maintain colony temperature and assure proper larval development (Heinrich, 1979a;O'Donnell and Foster, 2001), and acquire food for themselves and their colony (Heinrich, 1979a). Providing protection, care and food for the colony's young are the ways in which a non-reproductive forager increases its inclusive fitness.…”

Analysis of Hymenoptera Stings Reported to the Illinois Poison Center

“…Changes in air temperature and atmospheric pressure affect bee respiration and energy expenditure for ßight and hovering (Withers 1981, Burrill 1981) and also affect a beeÕs pollination activity. Furthermore, there have been studies that show that defensive behavior (stinging) is associated with both air temperature and wind speed (Drum and Rothenbuhler 1984, Southwick and Moritz 1987, Kastberger 2009). Human behavior also changes with weather conditions and seasonal holidays.…”

Subjective evaluation of defensive behavior in the Syrian honeybee (Apis mellifera syriaca)

“…32-16540 Hohen Neuendorf, Germany introduction Colony defense is a social behavior that involves a complicated sequence of actions by many individual bees and a variety of traits ranging from aggressive (e.g., guarding, recruiting, alerting, attracting, culminating, biting, stinging, and pursuing) to docile or gentle expression (Collins et al, 1980;Kastberger et al, 2009). An elaborate defensive system has evolved in both open-nesting (Seeley et al, 1982;Kastberger et al, 2008) and cavity-dwelling honeybee species (Ruttner, 1988), in tandem with the attractiveness of colonies and their nests as food resources for predators (Breed et al, 2004;Kastberger et al, 2009). In several species, defensive behavior was found to be heritable (Bell et al, 2009) and related to fitness (Dingemanse and Reale, 2005;Smith and Blumstein, 2008).…”

“…The docility of honeybee colonies is an important factor in practical beekeeping; consequently, this trait is an important selection trait in honeybee breeding (Bienefeld et al, 2007;. A variety of methods have been developed to evaluate defensive behavior in honeybee colonies, and assorted stimulation regimes have been used to measure behavior objectively (Stort, 1975;Woyke, 1992;Uribe-Rubio et al, 2008;Kastberger et al, 2009). However, several behavioral parameters cannot be measured easily and are, therefore, usually graded into classes according to observer judgment (Stevens, 1946;Wemelsfelder, 1997;Martin and Bateson, 1998;Wemelsfelder and Farish, 2004).…”