1. The underlying basis of alternative male reproductive strategies is either genetic or environmental. Several non-pollinating fig wasp lineages have dimorphic males, typically with winged males that disperse from natal figs to mate and flightless males that seek mating opportunities in natal figs.2. Walkerella sp. from Ficus benjamina has dark and pale wingless males. Observations and experiments in Xishuangbanna, Southern China found that (i) the sex ratio of Walkerella sp. did not vary with foundress number or brood size. (ii) The frequency of dark males increased with brood size and foundress number and they were absent from figs with a single foundress. This produced a higher proportion of dark males at higher densities. (iii) Males of both morphs fought, but injuries to dark males were more frequent. (iv) Dark males were more likely to disperse away from their natal figs and (v) they were more resistant to dehydration.3. Responses to selection are constrained by the genetic options available. Consequently, selection pressures acting on different lineages can produce similar outcomes that are achieved in different ways. Walkerella species lack winged males, but dark males display some of their features, dispersing from natal figs and displaying appropriate physiological and behavioural adaptations. However, dark males also displayed increased levels of damage from fighting -a feature unlikely to be shared with the winged males of other species.

Floral scents are known as an olfactory signal for attracting pollinators, but why the flowers pollinated by highly specialised pollinators emit scents consisting of mixtures of many compounds and dominated by one or a few compounds is still poorly understood. We supposed that each (especially characteristic) chemical in floral scents may play a specific role in mediating pollinator behaviours and tested this supposition in a fig-fig wasp mutualism. Ficus curtipes is obligately pollinated by an undescribed Eupristina species. In the scent of F. curtipes receptive figs, over 50 compounds have been identified, and the scent is dominated by two compounds, 6-methyl-5-hepten-2-ol (OL) and 6-methyl-5-hepten-2-one (NE). We therefore tested the roles of the two major chemicals in mediating the pollinator behaviours. Our results show that OL and NE, respectively, act as a longdistance attractant and a fig-entry behaviour stimulant to the obligate pollinator wasp. Namely, OL attracts the wasps to the figs and NE guides the wasps into the figs. This finding on the work division of floral scent compounds partially explains the maintenance mechanism of the fig-fig wasp mutualism and the significance of the chemical diversity of floral scent in plantpollinator interactions, especially in specialised pollination systems. ARTICLE HISTORY

The reproductive strategies of Walkerella sp.1 associated with Ficus curtipes and Walkerella sp.2 associated with Ficus benjamina were investigated. Both species oviposited from outside the fig wall. Walkerella sp.1 was the first non-pollinating fig to oviposit on Ficus curtipes and began to do this ten days after figs syconia began to develop. The larvae of Walkerella sp.1 were only found in the most external ovary layer of the fig. Walkerella sp.2 starts ovipositing after several other non-pollinating fig wasps have already laid their eggs in F. benjamina.The progeny of Walkerella sp.2 are distributed in the external ovary layer, the middle ovary layer, and/or the inner ovary layer of the figs. However, more than a quarter of the offspring were found in the most external layer and only a few in the inner layer. Experimental studies proved that the two Walkerella species are gall formers. In both manipulated figs and in natural figs, the sex ratios of Walkerella sp.1 and Walkerella sp.2 were female-biased. In Walkerella sp.2, the overall sex ratio increased with the proportion of figs parasitized in a crop, but this was not the case for Walkerella sp.1. Females of both Walkerella species appear not to have information about the patches on which they oviposit because sex ratios of both species decreased as brood sizes within individual figs increased and foundresses of both species were able to lay clutches containing a single male egg and several female eggs.