17Group living organisms rely on intra-group communication to adjust individual and 18 collective behavioural decisions. Complex communication systems are predominantly 19 multimodal and combine modulatory and information bearing signals. The honey bee 20 waggle dance, one of the most elaborate form of communication in invertebrates, 21 stimulates nestmates to search for food and communicates symbolic information 22 about the location of the food source. Previous studies on the dance behaviour in 23 diverse honey bee species demonstrated distinct differences in the combination of 24 visual, auditory, olfactory, and tactile signals produced by the dancer. We now studied 25 the behaviour of the receivers of the dance signals, the dance followers, to explore the 26 significance of the different signals in the communication process. In particular, we 27 ask whether there are differences in the behaviour of dance followers between the 3 28 major Asian honey bee species, A. florea, A. dorsata and A. cerana, and whether these 29 might correlate with the differences in the signals produced by the dancing foragers.
30Our comparison demonstrates that the behaviour of the dance followers is highly 31 conserved across all 3 species despite the differences in the dance signals. The highest 32 number of followers was present lateral to the dancer throughout the waggle run, and 33 the mean body orientation of the dance followers with respect to the waggle dancer 34 was close to 90° throughout the run for all 3 species. These findings suggest that dance 35 communication might be more conserved than implied by the differences in the 36 signals produced by the dancer. Along with studies in A. mellifera, our results indicate 37 that all honey bee species rely on tactile contacts between the dancer and follower to 38 communicate spatial information. The cues and signals that differ between the species 39 may be involved in attracting the followers towards the dancer in the different nest 40 environments. 41 42 Keywords 43 Apis florea, Apis cerana, Apis dorsata, waggle dance evolution, spatial information 44 transfer, tactile hypothesis 45 Cues and signals from the environment and nestmates can modulate the probability 72and intensity of dance behaviour [15,[17][18][19]. Interactions with nestmates in the hive 73 inform nectar foragers about the colony food stores and the nectar influx into the 74 colony [18,[20][21][22]. In addition, interactions with other foragers provide information 75 about predation and overcrowding at the food source [23][24][25][26]. An individual forager's 76 dance activity is modulated by the perceived reward value of the food source along 77 with information from these interactions [19]. This in turn drives recruitment to each 78 food source proportional to its relative reward value, which leads to an efficient 79 distribution of the colony's foraging force [27]. Thus, the waggle dance acts as the 80 primary regulatory mechanism of the colony's recruitment activity in addition to its 81 role in the effi...