Male-biased effective sex ratio across populations of the threatened Zamia boliviana (Zamiaceae)

Populations of dioecious plants commonly exhibit dissimilarities to the equilibrium expectation of a 1:1 sex ratio. Differential expenditure for reproduction between genders is cited as the primary mechanism responsible for a male-biased sex ratio, with increased effects on long-living species, but these mechanisms are still poorly understood. We explore the sex ratio in the endemic gymnosperm Zamia boliviana (Zamiaceae) populations from the Brazilian savanna (the Cerrado). We aim to investigate what the Z. boliviana sex ratio is, and whether population density and ecological correlates lead to variation in the sex proportion among Cerrado habitats. The study was conducted on ten in situ populations of Z. boliviana at sexual maturity (tertiary sex ratio). We estimated the populations’ sex ratio and performed a redundancy analysis to assess the relationship between biotic traits, such as sex ratio, and associated environmental features. Soil texture classes were used to classify the cycad habitats and were expressed in a ternary phase diagram. The results show a significant male-biased sex ratio in seven of the ten populations surveyed. Environmental factors did not explain the redundancy in the reproductive characteristics. However, the cycad occurs in different habitats in their endemic zone. Our study provides new biological data for Z. boliviana, suggesting that the differential reproductive expenditure of sexes in reproduction is governing the mechanisms of sex ratio variation, compared to local environmental factors in this cycad. The pattern of effective sex ratio found here improves our understanding of mechanisms causing biased sex ratios in cycads and other dioecious species from tropical ecosystems.