An Eltonian proxy for restoring a lost browser-tree interaction

Many South American dry woodlands lack good historical or paleoecological baseline data to inform restoration and conservation. However, functionalist approaches such as those popularized by rewilding suggest that functional interactions producing target ecosystem processes are valid even without data confirming compositionalist values such as a long coevolutionary histories or known historical range overlaps of target species. In central Chile, the guanaco (Lama guanicoe) has been extirpated but is known to browse trees in other regions of South America, and the tree Vachellia [Acacia] caven shows adaptations to browsing but has no extant browsers within its Chilean range. Both species are native to Chile but there are no data to assess their historical levels of interaction. Here we test the hypothesis that they can act as mutual "Eltonian proxy" species: interacting species for which we lack sufficient data (the Eltonian shortfall) to prove they are not proxies. Specifically we predict that they have complementary adaptations such that guanacos will browse Vachellia [Acacia] caven and the latter will show adaptive responses to their browsing. We introduced five guanacos into an enclosure of Vachellia [Acacia] caven "espinal" woodland, and over two years measured the growth responses of individual branches, compared to branches of trees in an area without browsing. We predicted that Vachellia [Acacia] caven would show compensatory growth in response to guanaco browsing resulting in an increase in branching. Guanacos browsed throughout the two years. In the presence of guanaco browsing, Vachellia [Acacia] caven branches grew longer, grew more sub-branches, and showed more densely streamlined branch architectures. These results indicate that guanacos could be used to substitute anthropogenic pruning as a restoration and management technique in Vachellia [Acacia] caven "espinal" woodlands. However, other extinct megaherbivores or extirpated deer may also be key components of a past herbivore community to which Vachellia [Acacia] caven was adapted. Further attention to a network of multiple interacting browsers, and their indirect and nontrophic effects, is an area for further research.