Variation in offspring sex ratios in the northern water snake (Nerodia sipedon)

We used data from 88 litters of northern water snakes (Nerodia sipedon) to test predictions about how mothers would adaptively vary the sex ratios of their offspring. Larger mothers produced significantly more daughters (r2 = 0.04, P = 0.05), and mothers producing larger offspring produced significantly more daughters (r2 = 0.06, P = 0.02). Because neonate size did not vary with maternal size, these sex-ratio patterns were independent of each other. These patterns were more pronounced for wild females than for females maintained in captivity while gravid, but rearing conditions did not have a significant effect on sex ratio. Also, because sex ratios were similar between captive and free-living females despite captive females giving birth 16 days earlier, on average, and because sex ratios did not vary with birth date within the two groups of females, gestation appeared not to affect sex ratio. If females vary sex ratios adaptively, only the relationship between sex ratio and neonate size was consistent with our predictions. Limited evidence from other snake species also indicates variation in neonatal sex ratios that is nonrandom but not necessarily adaptive. A better understanding of these patterns will require information on the factors that affect the fitness of male and female neonates differently. An unexpected sex-ratio pattern that we found was that 14 of 19 stillborn young were male. We speculate that this pattern could be a result of male embryonic sensitivity to temperature. Thus, the need for gravid females to maintain a high body temperature so that their young are born with enough time to find hibernation sites may conflict with the need for embryos to develop at a safe temperature.