Frugivory and seed dispersal by chelonians: a review and synthesis

In recent years, it has become clear that frugivory and seed dispersal (FSD) by turtles and tortoises is much more common than previously thought. We here review published and unpublished records of chelonian FSD, and assess the role of chelonians as seed dispersers, from individual species to the community level. We first discuss the distribution of chelonian FSD and the characteristics of the fruit and/or seed species eaten and dispersed by chelonians. We then use the seed dispersal efficiency framework to explore the quantitative and qualitative components of seed dispersal by tortoises and turtles, embarking on a journey from when the fruits and/or seeds are consumed, to when and where they are deposited, and assess how efficient chelonians are as seed dispersers. We finally discuss chelonian FSD in the context of communities and of chelonians as megafauna. A substantial proportion of the world's aquatic and terrestrial turtles and a major part of testudinid tortoises (71 species in 12 families) include fruits and/or seeds in their diet; fruits of at least 588 plant species in 121 families are ingested and/or dispersed by chelonians. For some chelonians, overall or in certain seasons, fruit may even form the largest part of their diet. Contrary to seed dispersal by lizards, the other major reptilian frugivores, chelonian FSD is not an island phenomenon in terms of geographic distribution. Nevertheless, on islands tortoises are often among the largest native terrestrial vertebrates - or were until humans arrived. We synthesise our knowledge of chelonian FSD, and discuss the relevance of our findings for conservation and restoration, especially in relation to rewilding with large and giant tortoises.