Sex- and Season-Dependent Behaviour in a Flightless Insect, the Auckland Tree Weta (Hemideina Thoracica)
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AvailableWehi et al.: on-line Seasonal at: http://www.newzealandecology.org/nzje/ behaviour patterns in weta 75 Sex- and season-dependent behaviour in a flightless insect, the Auckland tree weta (Hemideina thoracica) Priscilla M Wehi1,4*, Murray Jorgensen2 and Mary Morgan-Richards3 1Department of Biological Sciences, University of Waikato, Private Bag 3105, Hamilton 3240, New Zealand 2Department of Statistics, University of Waikato, Private Bag 3105, Hamilton 3240, New Zealand 3Institute of Natural Resources, Massey University, Private Bag 11222, Palmerston North, New Zealand 4Present address: CSAFE, University of Otago, P O Box 56, Dunedin, New Zealand *Author for correspondence (Email: [email protected]) Published online: 17 December 2012 Abstract: In a polygynous mating system, males frequently compete by locating and defending sites with resources essential to female survival and reproduction. We investigated seasonal changes in site occupancy in a sexually dimorphic, harem-forming insect, the Auckland tree weta (Hemideina thoracica). First we established artificial cavities as diurnal refuge cavities and potential harem guarding sites. We then examined cavity occupancy changes, and, based on our knowledge of prior occupants, determined sex-specific patterns of arrival, departure, and aggregation at a population level throughout the year. Both season and the sex of prior occupants influenced weta occupancy patterns. Most observations were of single females. However, both males and females moved into cavities previously occupied by a weta of the opposite sex more often than expected by chance alone. Females avoided cavities where other females were present, except during summer when most harems formed. In early summer, male and female tree weta previously living apart began co-habiting. Generally there was little relationship between the number and sex of the weta inside cavities and female departure rates from cavities. Males who were sharing with other males departed cavities more frequently than single males, as might be expected in a polygynous species with male–male combat. Males were less likely to depart if they were sharing a cavity with a harem of more than two females during the summer–autumn period. Analysis of departure rates from artificial cavities indicates males are more mobile than females only in winter and spring. Based on our arrival and departure data, and high occupancy of artificial cavities, we suggest that female weta at this site are mobile and may search for mates during the summer. The data are consistent with a polygynandrous mating system as inferred for other tree weta species (Hemideina spp.). Keywords: aggregation; harem; mating system; Orthoptera; polygyny Introduction Resource variation can lead to plasticity in mating systems so that, for example, mating system can vary within a species Searching for mates can be costly (Gwynne 1987; Acharya if resource availability varies spatially or temporally (Emlen 1995), but nonetheless is a requirement for most sexually & Oring 1977; Gwynne 1991). Seasonal effects can result reproducing organisms (Kokko & Wong 2007). Search effort in if resources vary throughout the year. Thus, for example, if one sex is dependent on search effort in the other (Hammerstein density is high, polygynous mating systems that have large & Parker 1987), but because males may benefit more from numbers of sneakers (or satellites), as well as residents, can multiple mating than females (Bateman 1948) males are favour mobile males (Reichard et al. 2004). Differences in generally predicted to mate-search more than females when aggregation and occupancy patterns between the sexes can multiple mating is common and sperm competition occurs thus occur both spatially and temporally as a consequence (Kokko & Wong 2007). However, males may attempt to of both sex-specific searching patterns and mating system monopolise resources, including females, if doing so increases (Lehmann & Perrin 2003). Nonetheless, data that report reproductive success. seasonal changes in sex-specific searching and aggregation In a polygynous mating system, males compete directly behaviour are predominantly from well-studied, large diurnal for access to females through male–male combat (cf. Darwin species. Data are still lacking for many taxa, and particularly 1874) or indirectly by locating and defending sites with for many insects. resources essential to female survival and reproduction (Emlen In many Orthopterans, males call, using stridulatory & Oring 1977; Shuster & Wade 2003; Kelly 2006a). If male ridges on their limbs or wings, and females search for mates, guarding of a limited resource (such as females, or refuge with females benefiting from male nuptial gifts such as cavities) increases reproductive success, then males might be spermatophores. However, sexually dimorphic New Zealand sedentary, and if females gain by enhancing their encounter tree weta (Hemideina spp., Anostostomatidae) have small rate with potential mates, they might be more mobile than spermatophores compared with other Orthoptera (Gwynne otherwise expected. The ability of individuals to monopolise 1995; Stringer 2001), and both males and females have matings will in turn influence the intensity of sexual selection stridulatory ridges. In six of the seven tree weta species, (Trivers 1972), particularly in polygynous species where mating males have enlarged mandibles used in male–male combat effort, and the opportunity for sexual selection for males, is (McIntyre 2001), and females form aggregations, or harems, high (Bateman 1948; Trivers 1985). in cavities used as day-time refugia (Field & Sandlant 2001). New Zealand Journal of Ecology (2013) 37(1): 75-83 © New Zealand Ecological Society. 76 New Zealand Journal of Ecology, Vol. 37, No. 1, 2013 These characteristics suggest that tree weta movement patterns tree saplings less than 2 m high. Predation pressure from associated with mating may not be typical of Orthopterans as introduced mammals and native owls was low (Morgan et al. a whole. If male guarding of a limited resource (e.g. females) 2009; PMW unpubl. data). Human disturbance was generally increases reproductive success, then males might be sedentary, low at night, when tree weta are active. Daytime disturbance, and if females gain by enhancing their encounter rate with however, was greater, with paths through the forest fragment potential mates, they might be more mobile than otherwise well used by members of the public. expected. However, there are few data on the formation or duration of aggregations, and of female harems that males Observations of movement patterns appear to defend (Field & Jarman 2001; Kelly 2006b). Recent research has highlighted potential negative effects of Tree weta are large-bodied (adults generally 3–6 g), tagging of study animals (e.g. Davis & Ovaska 2001; Boiteau flightless and nocturnal Orthopterans that seek refuge in tree et al. 2009; Barron et al. 2010; Saraux et al. 2011).Two trials cavities diurnally (Gibbs 1998). Eggs hatch in spring, and a to determine whether tagging might be a useful tool for this minimum of eight instars is required to reach adulthood in these study raised questions about the effects of tagging on tree weta hemimetabolous insects. Adults are long lived (adult lifespan movement. A majority of tree weta (93.4%; total n = 30) did not 9 months to 3 years; Leisnham et al. 2003). Because tree weta return to cavities after pronotum tagging with either bee tags or are nocturnal and their behaviour is difficult to observe directly, paper tags, and were not seen again either in these cavities or artificial cavities have proven useful to study tree weta ecology elsewhere, despite data suggesting that Hemideina thoracica and behaviour (e.g. Trewick & Morgan-Richards 1995). and H. crassidens individuals most likely have high site fidelity Hypotheses about the genus have developed in large part from (Spurr & Berben 2004; PMW unpubl. data). Given this result, work on Hemideina crassidens Blanchard 1851 (e.g. Moller and the results from a previous tagging study (Ordish 1992), 1985; Kelly 2006a, b) and Hemideina maori Pictet & Saussure we minimised disruption by recording data from unmarked 1891 (e.g. Jamieson et al. 2000; Leisnham & Jamieson 2002; individuals; individual tracking is not essential to understand Leisnham et al. 2003). In contrast, little is known of Auckland occupancy and movement patterns at the population level. tree weta (Hemideina thoracica White 1846) ecology despite its extensive distribution throughout much of the North Island, Refuge design and monitoring New Zealand. Field studies of Hemideina crassidens indicate Forty artificial refuges, consisting of two cavities each, were that females apparently prefer unoccupied cavities, males constructed from aged Podocarpus totara (tōtara) timbers defend females not cavities, and males are more transient and attached to 20 randomly selected, mature Dacrycarpus than females (Kelly 2006c). Nonetheless, most of these field dacrydioides trees (Fig. 1). Refuges were attached vertically studies were undertaken in weta populations where more refuge at 2.5 m and their direction randomised on each tree. Refuge cavities were empty than were occupied, in young forest (less design was generally similar to that suggested in Trewick & than 30 years old), and with observations undertaken during Morgan-Richards (2000) and Spurr & Berben (2004), but on a the late summer and autumn only. We investigated occupancy patterns