The rate of biodiversity loss is not slowing despite global commitments, and the depletion of animal species can reduce the stability of ecological communities. Despite this continued loss, some substantial progress in reversing defaunation is being achieved through the intentional movement of animals to restore populations. We review the full spectrum of conservation translocations, from reinforcement and reintroduction to controversial conservation introductions that seek to restore populations outside their indigenous range or to introduce ecological replacements for extinct forms. We place the popular, but misunderstood, concept of rewilding within this framework and consider the future role of new technical developments such as de-extinction.

Starting in the late 1970s, ecologists began unraveling the role of recently extinct large vertebrates in evolutionary ecology and ecosystem dynamics. Three decades later, practitioners are now considering the role of ecological history in conservation practice, and some have called for restoring missing ecological functions and evolutionary potential using taxon substitutes Á extant, functionally similar taxa Á to replace extinct species. This pro-active approach to biodiversity conservation has proved controversial. Yet, rewilding with taxon substitutes, or ecological analogues, is now being integrated into conservation and restoration programmes around the world. Empirical evidence is emerging that illustrates how taxon substitutions can restore missing ecological functions and evolutionary potential. However, a major roadblock to a broader evaluation and application of taxon substitution is the lack of practical guidelines within which they should be conducted. While the International Union for Conservation of Nature's reintroduction guidelines are an obvious choice, they are unsuitable in their current form. We recommend necessary amendments to these guidelines to explicitly address taxon substitutions. A second impediment to empirical evaluations of rewilding with taxon substitutions is the sheer scale of some proposed projects; the majority involves large mammals over large areas. We present and discuss evidence that large and giant tortoises (family Testudinidae) are a useful model to rapidly provide empirical assessments of the use of taxon substitutes on a comparatively smaller scale. Worldwide, at least 36 species of large and giant tortoises went extinct since the late Pleistocene, leaving 32 extant species. We examine the latent conservation potential, benefits, and risks of using tortoise taxon substitutes as a strategy for restoring dysfunctional ecosystems. We highlight how, especially on islands, conservation practitioners are starting to employ extant large tortoises in ecosystems to replace extinct tortoises that once played keystone roles.

There is increasing evidence that restoration ecologists should be most concerned with restoring species interactions rather than species diversity per se [1]. Rewilding with taxon substitutes, the intentional introduction of exotic species to replace the ecosystem functions of recently extinct species, is one way to reverse ecosystem dysfunction following the loss of species interactions [2]. This is highly controversial [3], in part because of a lack of rigorous scientific studies [4]. Here we present the first empirical evidence of an in situ rewilding project undertaken as a hypothesis-driven ecosystem management option. On Ile aux Aigrettes, a 25-hectare island off Mauritius, the critically endangered large-fruited endemic ebony, Diospyros egrettarum (Ebenaceae), was seed-dispersal limited after the extinction of all native large-bodied frugivores, including giant tortoises. We introduced exotic Aldabra giant tortoises, Aldabrachelys gigantea, to disperse the ebony seeds. Not only did the tortoises ingest the large fruits and disperse substantial numbers of ebony seeds, but tortoise gut passage also improved seed germination, leading to the widespread, successful establishment of new ebony seedlings. Our results demonstrate that the introduction of these exotic frugivores is aiding the recovery of ebonies. We argue for more reversible rewilding experiments to investigate whether extinct species interactions can be restored.