Exploring the reproductive ecology of the tropical semifossorial snake  Ninia atrata

Based on histological analyses and field studies, this research describes the reproductive ecology of a population of Ninia atrata snakes inhabiting an oil palm plantation. Furthermore, through a multivariate approach, we explored the main drivers of reproductive output in N. atrata. Results showed that prey abundance and food intake were crucial variables contributing to reproductive output. Multiple linear regression models showed that neonates had high sensitivity (R²=55.29%) to extreme changes in climate, which was strongly related to slug and snail abundance variability and microhabitat quality. Reproductive cycles were markedly different between the sexes, being continuous in males and cyclical in females. Despite this variation, reproductive cycles at the population level were seasonal semi-synchronous. Constant recruitment of neonates all year, multiple clutches, high mating frequency, and continuous sperm production characterized the reproductive phenology of N. atrata. In addition, a significant number of previtellogenic females presented oviductal sperm as well as uterine scars, suggesting a high precocity in the species. The main drivers of reproductive output also differed between the sexes. In females, clutch size and secondary follicle variability were highly related to stomach bolus volume, fat body area, and body mass. In males, height of piles of palm leaves and body mass, rather than intrinsic reproductive traits, were the main drivers of sperm production. Nevertheless, in both cases, the relationship between body mass, prey abundance, and food intake suggests that N. atrata follows the income breeding strategy to compensate for reproductive costs and to maximize fitness.

Keywords
Continuous male reproduction; Clutch mass; Income breeding; Iteroparity; Spermatogenesis; Oogenesis; Reproductive effort; El Niño-Southern Oscillation (ENSO)