Communication in the Hymenoptera Chemistry, Ecology and Evolution Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universität Würzburg vorgelegt von Thomas Schmitt Würzburg Würzburg, 2004 Eingereicht am: ............................................................................. Mitglieder der Promotionskommission: Vorsitzender: ................................................................................ 1. Gutachter: Prof. Dr. Erhard Strohm 2. Gutachter: Prof. Dr. Peter Schreier Tag des Promotionskolloquiums: ................................................. Doktorurkunde ausgehändigt am: ................................................ This thesis is based on the following manuscripts: 1. Schmitt T., Herzner G., Schreier P., Strohm E.: Volatiles of foraging honeybees Apis mellifera L. (Hymenoptera: Apidae) and their possible role as semiochemicals and kairomones. In prep. (Chapter 2) 2. Herzner G., Schmitt T., Linsenmair K. E. and Strohm E.: Prey recognition by females of the European beewolf and its potential for a sensory trap. In prep. (Chapter 3) 3. Schmitt T., Strohm E., Herzner G., Bicchi C., Krammer G., Heckel F., Schreier P. (2003): (S)-2,3-Dihydrofarnesoic acid, a new component in cephalic glands of male European beewolves Philanthus triangulum. Journal of Chemical Ecology 29(11): 2469-2479 (Chapter 4) 4. Herzner G., Schmitt T., Schreier P., Strohm E.: Brothers smell similar: Sex pheromone variation in a wasp and its implications for inbreeding avoidance. In prep. (Chapter 5) 5. Strohm E., Schmitt T., Herzner G., Laurien-Kehnen C., Boland W., Schreier P.: A cuckoo in wolves’ clothing? Chemical mimicry in a specialised cuckoo wasp of the European beewolf (Hymenoptera, Chrysididae and Crabronidae). In prep. (Chapter 6) 6. Endler A., Liebig J., Schmitt T., Parker J., Jones G., Schreier P., Hölldobler B. (2004): Surface hydrocarbons of queen eggs regulate worker reproduction in a social insect. Proceedings of the National Academy of Science of the USA 101(9): 2945-2950 (Chapter 7) 7. Brandt M., Heinze J., Schmitt T., Foitzik S. (in press): A chemical level in the coevolutionary arms race between an ant social parasite and its hosts. Journal of Evolutionary Biology (Chapter 8) 1 Contents Contents............................................................................................................................1 1 General Introduction......................................................................................................3 2 Volatiles of foraging honeybees Apis mellifera L. (Hymenoptera: Apidae) and their possible role as semiochemicals and kairomones .....................................13 3 Prey recognition by females of the European Beewolf and its potential for a sensory trap............................................................................................................25 4 (S)-2,3-Dihydrofarnesoic acid, a new component in cephalic glands of male European beewolf Philanthus triangulum........................................................45 5 Brothers smell similar: Sex pheromone variation in a wasp and implications for inbreeding avoidance .....................................................................59 6 A cuckoo in wolves’ clothing? Chemical mimicry in a specialised cuckoo wasp of the European beewolf (Hymenoptera, Chrysididae and Crabronidae) .............................................................................................................81 7 Surface hydrocarbons of queen eggs regulate worker reproduction in a social insect ............................................................................................................101 8 A chemical level in the coevolutionary arms race between an ant social parasite and its hosts...............................................................................................117 9 General Discussion....................................................................................................141 10 Summary..................................................................................................................149 11 Zusammenfassung ...................................................................................................151 12 Curriculum Vitae .....................................................................................................155 13 Danksagung .............................................................................................................159 2 3 1 General Introduction 1.1 Communication All social interactions between organisms, be it courtship, species and kin recognition, division of labour or any other form of coexistence, involve communication. Animal signals evolved in a diversity of forms constrained by the environment in which the signal is propagated and the physiological equipment of sender and receiver. This generated different sensory modes of communication: chemical, auditory, visual and tactile communication. (Dusenbery 1992; Bradbury & Vehrencamp 1998; Greenfield 2002). However, how a signal in a certain modality evolved its characteristics is not known in most cases. This study contributes to the understanding of the evolution of communication. But what is communication? To understand the evolution of communication it is necessary to define the different types of communication. However, the definition of communication is still under debate and definitions given by leading scientists in the field of animal communication studies vary considerably (Hölldobler 1984). Wilson (1975) defined biological communication as “action on the part of one organism (or cell) that alters the probability pattern of behaviour in another organism (or cell) in an adaptive fashion. By adaptive I mean that the signalling, or the response, or both have been genetically programmed to some extent by natural selection”. This definition is more general than ones, which limit “true communication” to situations where the signal is generated by the sender to provide information and the response is adaptive for the receiver (e. g. Dusenbery 1992, p. 37). Dusenbery’s definition would confine “true communication” to signals within a species, beneficial to the receiver. However, taking into account the conflicts even between the sexes within a species, communication that is equally adaptive for both signaller and receiver is likely to be rare. Dawkins (1995), in contrast, gave a more simple definition: “communication occurs when one animal’s 4 behaviour can be shown to have an effect on the behaviour of another. “Signals” are the means by which these effects are achieved” (Dawkins 1995, p. 73). This shows that communication is one of the most contentious issues in animal behaviour and there is no commonly accepted definition. However, nearly all authors agree that communication involves the provision of information (cues or signals) by a sender, and the subsequent use of that information by a receiver in deciding how to respond (Bradbury & Vehrencamp 1998, p. 2). This definition allows to include deception by the sender and eavesdropping or other kinds of signal exploitation by the receiver. It is therefore useful to distinguish between cases of cooperative signalling, in which both, sender and receiver, benefit and of non-cooperative signalling, in which only the sender or only the receiver benefits (Harper 1991; Johnston 1997). 1.2 Signal Evolution To understand the evolution of communication, one has to consider two sets of selective pressures. First, selective pressures acting on signallers, second, selective pressures acting on receivers. Thus, natural selection favour signallers to be more effective in manipulating the receiver and having lower fitness costs by, for example, choosing less costly signals or avoid eavesdropping (Wiley 1983, 1984; Endler 1992, 1993). On the other hand, selection favours receivers who can adequately adjust their behaviour in response to the information provided by signals (Johnston 1997). Besides natural selection one has to consider other potentially important nonadaptive forces such as developmental and phylogenetic constraints, pleiotropy and genetic drift (Phelan 1992). Yet another important evolutionary factor are preadaptations, which evolve in other contexts than communication and signal design. Scenarios for signal origin also fall into two categories: sender preadaptations and receiver preadaptations. The classical ethological view is that signals evolve from sender preadaptations such as behavioural, physiological, or morphological traits (Otte 1974). If the presence of the trait provides 5 useful information to receivers that are able to detect it, receivers will evolve to pay attention to the trait and use it to make behavioural decisions in response. Signals that evolve from this type of source contain information from the very beginning because the sender’s incipient signal is linked to its condition. Both, sender and receiver might gain benefits from this type of communication. However, considering the conflict that exists between sender and receiver in the majority of cases, even in a cooperative signalling mode, it is conceivable that either the sender or receiver gains larger or exclusive benefits (Smith & Harper 2003). Thus, cues or signals can also evolve to deceit the receiver or to manipulate the receiver’s response. From the signaller’s perspective, a signal is a means of manipulation. It serves to influence the receiver’s behaviour in a way that benefits the signaller.