Environmental variation structures reproduction and recruitment in long-lived mega-herbivores: Galapagos giant tortoises

Migratory, long-lived animals are an important focus for life-history theory because they manifest extreme trade-offs in life-history traits: delayed maturity, low fecundity, variable recruitment rates, long generation times, and vital rates that respond to variation across environments. Galapagos tortoises are an iconic example: they are long-lived, migrate seasonally, face multiple anthropogenic threats, and have cryptic early life-history stages for which vital rates are unknown. From 2012 to 2021, we studied the reproductive ecology of two species of Galapagos tortoises (Chelonoidis porteri and C. donfaustoi) along elevation gradients that coincided with substantial changes in climate and vegetation productivity. Specifically, we (1) measured the body and reproductive condition of 166 adult females, (2) tracked the movements of 33 adult females using global positioning system telemetry, and monitored their body condition seasonally, (3) recorded nest temperatures, clutch characteristics, and egg survival from 107 nests, and (4) used radiotelemetry to monitor growth, survival, and movements of 104 hatchlings. We also monitored temperature and rainfall from field sites, and remotely sensed primary productivity along the elevation gradient. Our study showed that environmental variability, mediated by elevation, influenced vital rates of giant tortoises, specifically egg production by adult females and juvenile recruitment. Adult females were either elevational migrants or year-round lowland residents. Migrants had higher body condition than residents, and body condition was positively correlated with the probability of being gravid. Nests occurred in the hottest, driest parts of the tortoise's range, between 6 and 165 m elevation. Clutch size increased with elevation, whereas egg survival decreased. Hatchling survival and growth were highest at intermediate elevations. Hatchlings dispersed rapidly to 100-750 m from their nests before becoming sedentary (ranging over <0.2 ha). Predicted future climates may impact the relationships between elevation and vital rates of Galapagos tortoises and other species living across elevation gradients. Resilience will be maximized by ensuring the connectivity of foraging and reproductive areas within the current and possible future elevational ranges of these species.