Relictness: A Relative Notion and the Need for Formal Analyses
The term relict is generally employed either for emblematic taxa or for well-defined situations where taxonomic, phylogenetic and paleontological characterizations are established from previous studies and publications. This definition generally embodies the relict species, its very large sister-group, the paleontological record and the geographic restriction if any. For example, the relict Amborella trichopoda is the sister-group of all other flowering plants and it is only found in New Caledonia (Soltis et al. 2002). But relictness is not a particular state of a character that can be distinguished unambiguously from other and different states. Like most other classlevel characterizations such as rarity, specialization or endemism (e.g., Rabinowitz 1981; Futuyma and Moreno 1988; Anderson 1994), relictness is a relative and composite situation that needs to be established by comparison, and in every case by phylogenetic comparison. Strictly speaking, we should not say Amborella trichopoda is “a relict” but Amborella trichopoda is “a relict species for flowering plants.” This comparison is based on topology and branch lengths that depend on the taxon and character samples used to build the tree. This way, within flowering plants, there are many groups that stand isolated on long branches and could be called relicts, such as Welwitschia, Ephedra, etc., actually hundred of species among hundreds of thousands of plant lineages (for examples see Jacobson and Lester 2003; Dilcher et al. 2005). As it is set by comparison between sister-groups within a phylogenetic tree, characterization as a relict will depend on the taxon sample used in that tree. For non-phylogeneticists, this could sound like a limitation of this notion that makes it less useful. Actually, a statement of relictness needs to be based on a formal phylogenetic analysis conducted on a given set of taxa. Depending on the tree obtained, a gap analysis can show that one or several branches have exceptional lengths and originate deep in the tree. These branches and their terminal taxa can be named relict taxa. This is required to implement the phylogenetic diversity criterion for conservation, characterizing the extreme case of relicts at the same time.
Relicts and Ecosystem Functioning
Macroevolutionary studies of this type might appear to be far removed from the real nowaday's world where ecosystems must function and populations must be viable to be conserved. Actually, historical and functional views are not opposed or disconnected (Brooks and McLennan 1991; Grandcolas 1998). Current research (e.g., Elias et al. 2013) in the framework of community phylogenetics (Ricklefs and Latham 1992; Webb et al. 2002) shows that trophic webs have a phylogenetic structure. Phylogenetic niche conservatism mitigated by exploitative competition means that related species can have similar resource use (Cadotte et al. 2008; but see Mouquet et al. 2012). In this theoretical framework, a relict is then expected to exploit a unique niche, a prediction consistent with some of the adaptive explanations cited above (e.g., Parsons 2005), that relicts can be highly specialized (but inconsistent with relicts as generalists).
Therefore, maximizing phylogenetic diversity for conservation can be expected to select for species whose resource use is unique (Srivastava et al. 2012; Winter et al. 2013). In cases where relicts are found in a very stable and specialized habitat harboring small communities, this original resource use might implicate a key ecosystem service (e.g., Gibert and Deharveng 2002). At the extreme, structuring ecological communities by conserving species on the basis of phylogenetic diversity should select against loss of function in communities, by retaining species with lower niche overlap even if ecological redundancy is decreased.