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The existence of spiteful behaviors remains controversial. Spiteful behaviors are those that are harmful to both the actor and the recipient, and they represent one of the four fundamental types of social behavior (alongside selfishness, altruism, and mutual benefit). It has generally been assumed that the conditions required for spite to evolve are too restrictive, and so spite is unlikely to be important. This idea has been challenged in recent years, with the realization that localized competition can relax the conditions required for spite to evolve. Here we develop a theoretical model for a prime candidate for a spiteful behavior, the production of the sterile soldier caste in polyembryonic wasps. Our results show that (a) the biology of these soldiers is consistent with their main role being to mediate conflict over the sex ratio and not to defend against competitors and (b) greater conflict will occur in more outbred populations. We also show that the production of the sterile soldier caste can be classed as a spiteful behavior but that, to an extent, this is merely a semantic choice, and other interpretations such as altruism or indirect altruism are valid. However, the spite interpretation is useful in that it can lead to a more natural interpretation of relatedness and facilitate the classification of behaviors in a way that emphasizes biologically interesting differences that can be empirically tested.

Oil palm plantations cover %14.6 million ha worldwide and the total area under cultivation is expected to increase during the 21st century . Indonesia and Malaysia together account for 87% of global palm oil production and the combined harvested area in these countries has expanded by 6.5 million ha since 1990. Despite this, soil C cycling in oil palm systems is not well quantified but such information is needed for C budget inventories. We quantified soil C storage (root biomass, soil organic matter (SOM) and microbial biomass) and losses [potential soil respiration (R s ) and soil surface CO 2 flux (F s )] in mineral soils from an oil palm plantation chronosequence (11-34 years since planting) in Selangor, Malaysia. There were no significant effects of plantation age on SOM, microbial biomass, R s or F s , implying soil C was in dynamic equilibrium over the chronosequence. However, there was a significant increase in root biomass with plantation age, indicating a short-term C sink. Across the chronosequence, R s was driven by soil moisture, soil particle size, root biomass and soil microbial biomass N but not microbial biomass C. This suggests that the nutrient status of the microbial community may be of equal or greater importance for soil CO 2 losses than substrate availability and also raises particular concerns regarding the addition of nitrogenous fertilizer, i.e. increased yields will be associated with increased soil CO 2 emissions. To fully assess the impact of oil palm plantations on soil C storage, initial soil C losses following land conversion (e.g. from native forest or other previous plantations) must be accounted for. If initial soil C losses are large, our data show that there is no accumulation of stable C in the soil as the plantation matures and hence the conversion to oil palm would probably represent a net loss of soil C.