Global energy gradients and size in colonial organisms: Worker mass and worker number in ant colonies

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Abstract
Body mass shapes processes from cell metabolism to community dynamics. Little is known, however, about how the average body mass of individuals varies among ecological communities. Ants alter colony mass by independently changing worker mass and/or worker number. In a survey of 49 ecosystems from tundra to tropical rainforest, average worker mass and worker number were uncorrelated (r s = 0.2, P > 0.14) and varied 100-fold. Data supported the hypothesis that higher mean monthly temperatures, T, reduce worker mass by increasing metabolic costs during worker development. In contrast, worker number was unimodal over a 1,000-fold gradient of net primary productivity (NPP, g of carbon per m2 per yr), a measure of organic carbon available to consumers. At the lowest levels of NPP colonies appeared to be carbon-limited; above 60 g of carbon per m2 per yr average worker number decreased to a global low. This decline in worker number with increasing NPP supports the hypothesis that abundant carbon ameliorates the Achilles heel of small taxa in competition with large taxa: their relatively high metabolic demands. Higher predation rates in resource-rich environments may also play a role in limiting worker number. In all, about half the global variation in worker mass and number was accounted for by gradients of NPP and T. Changes in global temperature and rainfall may thus mold gradients of ectotherm size, with consequences for the structure and function of the ecosystems.