Feminization of a mixed-stock foraging aggregation of immature green turtles (Chelonia mydas), 1975-2018

Marine turtles exhibit temperature-dependent sex determination (TSD), and there is widespread concern that global warming is raising nest incubation temperatures, resulting in increasingly female-skewed sex ratios in "feminized" populations. We assessed the sex ratio of a mixed-stock aggregation of immature green turtles (Chelonia mydas) at a midocean developmental foraging ground in the Northwest Atlantic from 1975 to 2018. We used plasma testosterone concentrations, measured by radioimmunoassay (RIA) and calibrated by laparoscopy, to determine the sex of 2,724 green turtles captured 3,940 times in Bermuda (32 degrees 18 ' N, 64 degrees 46 ' W) waters. A logistic regression model correctly predicted the sex of 99.5% of turtles (189/190) with associated testosterone concentrations whose sex had been verified via laparoscopy. Empirical evaluation of the trend in sex ratio using four related data sets showed a significant increase (2.8-4.0% yr(-1)) in the percent females, ranging from 62.7 to 68.1% during the most recent years of evaluation. Using growth rates to predict the year of arrival of turtles in Bermuda, we estimated the sex ratio of recruiting cohorts over 4 decades. Mixed-stock analysis of mtDNA sequences of 602 turtles that recruited to Bermuda between 1970 and 2018 suggested that multiple, geographically dispersed rookeries contributed to the Bermuda aggregation making it regionally representative. Changes in rookery contributions and strong population increases at certain rookeries may partly explain the increasing trend in the percent female. But the steady rate of increase over decades and the increasing female percentage of arriving cohorts are consistent with impacts of global warming at source rookeries.