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Ewers & Cowley FINAL.P65 The role of sound production in determining dominance in agonistic interactions between male tree wetas (Hemideina crassidens, Orthoptera: Anostostomatidae) Robert M. Ewers1,2,3 and Greig Cowley1 1 School of Biological Sciences, University of Canterbury, Private Bag 4800, Christchurch, New Zealand 2 Current address: Institute of Zoology, Zoological Society of London, Regents Park, London NW1 4RY, UK, and Department of Zoology, Cambridge University, Downing Street, Cambridge CB2 3EJ, UK 3Corresponding author’s email: [email protected] (Received 17 July 2005, revised and accepted 5 October 2005) Abstract New Zealand tree wetas in the genus Hemideina have a resource-defence polygynous mating system, where males compete for ownership of galleries and the females they contain. Large males have previously been shown to have better success in male-male agonistic encounters than small males, and consequently obtain more mating opportunities. However, sound production by stridulation is frequently observed in agonistic interaction between males, but its role in determining the outcome is unknown. Here, we experimentally tested the role of sound production in determining dominance in the tree weta H. crassidens. We found that muting a dominant male can result in a change in dominance between rival males, but only if the males are similar in size. Large disparities in the size of combating males overrides any effect of stridulation. Keywords: agonistic interaction - body size - dominance - Hemideina - stridulation - sound production - weta. Introduction frequently found in the lowland forests of Westland, Nelson and Marlborough in New Zealand tree wetas (Hemideina spp.) the South Island, and are also common are endemic, flightless orthopterans, and in Wellington in the North Island (Field are among the heaviest insects in the & Rind 1992). Hemideina wetas exhibit world (Field 2001b, Koning & Jamieson strong sexual dimorphism, with females 2001). Of this genus, H. crassidens is typically having larger body size than one of the most common in the South males (Kelly 2005), but males having Island (Field & Rind 1992). They are greatly enlarged heads and mandibles (Field New Zealand Natural Sciences (2005) 30: 11-17 12 New Zealand Natural Sciences 30 (2005) & Sandlant 1983, Kelly 2005) which they which has been observed both during use to fight for possession of galleries. (Field 1993a) and after (Field & Rind Galleries inhabited by tree wetas are most 1992) combat between males (Field typically small, tunnel-like chambers in 2001a). Stridulation in agonistic trees (Ordish 1992), but for the alpine encounters can be used by both an wetas, H. maori, are cavities between rocks intruding weta, to signal aggressive intent, (Jamieson 2002). Regardless of the gallery and by the resident weta, to signal type, male wetas compete with each other resistance to the intruder (Field 2001a). for access to galleries and the females that As such, it is likely that stridulation is shelter in them during the day (Field & used as form of threat display that is Sandlant 1983, 2001, Koning & Jamieson analagous to visual displays of mandible 2001, Kelly 2005). This behaviour size (Field 1993a). If stridulation is a involves an intruder male attempting to strong signal of fighting ability, it is remove a resident male from the gallery, possible that preventing a weta from thereby simultaneously taking possession stridulating could result in that weta of the gallery and the female(s) within. losing dominance in an agonistic This type of interaction is stereotypical encounter. Here, we tested the role of of a resource defence polygynous mating sound production in determining the system (Field & Sandlant 1983, Koning establishment of dominance between male & Jamieson 2001). wetas by experimentally muting males. It is already been well established that larger males are more successful in gaining Materials and methods control of galleries and mates (Gwynne & Jamieson 1998, Field 2001a, Koning Nine male and nine female adult wetas & Jamieson 2001, Jamieson 2002, were collected from native forest near Leisnham & Jamieson 2004, Kelly 2005), Hari-Hari (West Coast, South Island). although surprisingly, this does not Just nine males were collected as the necessarily equate to higher copulation collection methods are destructive and success (Leisnham & Jamieson 2004). result in the complete demolition of However, one aspect of male-male galleries in the habitat (Field & Sandlant competition that has received little 2001). Male-female pairs were randomly attention is the role of sound production selected and placed into cylindrical plastic during agonistic interactions. Sound is containers (20 cm diameter, 25 cm deep) produced from a stridulatory apparatus where they were fed with raw carrots and comprised of hundreds of small pegs on fresh Pittosporum sp. leaves, and water was the femora which are rubbed against files provided in a vial plugged with cotton on abdominal tergite II (Ordish 1992, wool (after Field 1993a). They were kept Field 1993a, 1993b). This appears to be at 15°C on a reverse light cycle that the principal means of communication alternated 12 hours of light with 12 hours amongst individuals (Ordish 1992) and of darkness. Because wetas are nocturnally occurs in three behavioural contexts; active, experiments were conducted defence, territorial signalling and agonistic during the night phase of the cycle. interactions between rival males (Ordish Observations were made under red-light 1992, Field 1993a). Of interest here is and also recorded using infra-red video stridulation in agonistic encounters, for later analysis. Before any experiments R.M. EWERS & G. COWLEY: Stridulation in weta agonistic interactions 13 were conducted, all carrot, leaves and water status in the re-match as they did in the vials were removed from the container. original trial. Ten pairs of male combatants were One possible confounding effect that randomly selected, with each of the nine could influence the outcome of re-matches males confronting several different between males is that individuals may opponents (range 1 – 3). Male-male trials remember the outcome of previous were initiated by introducing a male into interactions (Field & Rind 1992). Data a container that already contained a male- on the temporal extent of memory in female pair. Observations continued until Hemideina weta are unavailable, but one male had proven dominance over the studies on the field cricket Gryllus other by copulating with the female. bimaculatus (Orthoptera: Gryllidae) The male introduced to initiate a trial indicate that memory of agonistic was called the intruder, and the male interactions in Orthopterans lasts for less already present was the resident. All males than 24 hours (Adamo & Hoy 1995, were weighed on an electronic balance to Khazraïe & Campan 1997, 1999). To the nearest 0.01 g, and head width, minimise the likelihood that memory mandible length, and hind leg tibia length played a role in our experiment, muted were measured with calipers to the nearest individuals were left undisturbed for at 0.1 mm. Relationships between the body least four days before being re-matched attributes were assessed with correlation with the previously subordinate male. analysis, and individuals were assigned The role of muting in determining two trait-based index values. First, all changes in dominance was tested with a individuals were given a score (S) for each Chi-squared goodness-of-fit test (Zar attribute (A), calculated by dividing the 1996). The way in which body size value for that individual by the maximum influenced the outcome of this experiment value obtained for that attribute from all was tested with a binomial general linear the measured wetas (maxSA). This is model (GLM), in which the difference in represented by the formula relative body size of the two combatants was used to predict the probability of a change in dominance following muting. where SAi is the score S for body attribute A in individual i. Scores for each of the Results four attributes were averaged and squared to give an overall index of relative body Four residents and six intruders established size (BI) for the nine wetas with the dominance following agonistic formula interactions, and all body attributes were positively correlated with all others (Table 1). The index of body size BI was a strong The dominant male from each trial was predictor of dominance, with larger males muted by applying a low melting point more likely to dominate agonistic wax (75 % resin and 25 % beeswax) to encounters than small males (linear the stridulatory ridges on the hind legs regression on log10-transformed BI; and abdomen. The ten male-male pairs F1,7 = 7.35, P = 0.03, r = 0.72; Figure 1). were then re-matched in a second trial. The application of wax to a male’s The males had the same occupational stridulatory ridges prevented the 14 New Zealand Natural Sciences 30 (2005) Table 1. Results of Pearson correlation analysis between body attributes of male wetas. Correlation rn P head width - mandible length 0.98 9 <0.001 head width - hind tibia length 0.89 9 0.001 head width - mass 0.86 9 0.003 mandible length - hind tibia length 0.91 9 0.001 mandible length - mass 0.82 9 0.006 tibia length - mass 0.80 9 0.009 Figure 1. Relationship between body size of male wetas and the probability of them winning agonistic encounters. Body size is an index based on measurements of head width, mandible length, hind tibia length and body mass (see text
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