Demographic outcomes and ecosystem implications of giant tortoise reintroduction to Española Island, Galapagos.

Restoration of extirpated species via captive breeding has typically relied on population viability as the primary criterion for evaluating success. This criterion is inadequate when species reintroduction is undertaken to restore ecological functions and interactions. Herein we report on the demographic and ecological outcomes of a five-decade-long population restoration program for a critically endangered species of "ecosystem engineer": the endemic Española giant Galapagos tortoise (Chelonoidis hoodensis). Our analysis of complementary datasets on tortoise demography and movement, tortoise-plant interactions and Española Island's vegetation history indicated that the repatriated tortoise population is secure from a strictly demographic perspective: about half of tortoises released on the island since 1975 were still alive in 2007, in situ reproduction is now significant, and future extinction risk is low with or without continued repatriation. Declining survival rates, somatic growth rates, and body condition of repatriates suggests, however, that resources for continued population growth are increasingly limited. Soil stable carbon isotope analyses indicated a pronounced shift toward woody plants in the recent history of the island's plant community, likely a legacy of changes in competitive relations between woody and herbaceous plants induced by now-eradicated feral goats and prolonged absence of tortoises. Woody plants are of concern because they block tortoise movement and hinder recruitment of cactus--a critical resource for tortoises. Tortoises restrict themselves to remnant cactus patches and areas of low woody plant density in the center of the island despite an apparent capacity to colonize a far greater range, likely because of a lack of cactus elsewhere on the island. We conclude that ecosystem-level criteria for success of species reintroduction efforts take much longer to achieve than population-level criteria; moreover, reinstatement of endangered species as fully functioning ecosystem engineers may often require large-scale habitat restoration efforts in concert with population restoration.