Plant cooperation.

The study of plant behaviour will be aided by conceptual approaches and terminology for cooperation, altruism and helping. The plant literature has a rich discussion of helping between species while the animal literature has an extensive and somewhat contentious discussion of within-species helping. Here, I identify and synthesize concepts, terminology and some practical methodology for speaking about helping in plant populations and measuring the costs and benefits. I use Lehmann and Keller's (2006) classification scheme for animal helping and McIntire and Fajardo's (2014) synthesis of facilitation to provide starting points for classifying the mechanisms of how and why organisms help each other. Contextual theory is discussed as a mechanism for understanding and measuring the fitness consequences of helping. I synthesize helping into four categories. The act of helping can be costly to the helper. If the helper gains indirect fitness by helping relatives but loses direct fitness, this is altruism, and it only occurs within species. Helpers can exchange costly help, which is called mutualism when between species, and reciprocation when within a species. The act of helping can directly benefit the helper as well as the recipient, either as an epiphenomenon resulting from behaviours under natural selection for other reasons, or because the helper is creating a mutual benefit, such as satiating predators or supporting a mutualism. Facilitation between species by stress amelioration, creation of novel ecosystems and habitat complexity often meets the definition of epiphenomenon helping. Within species, this kind of helping is called by-product mutualism. If the helping is under selection to create a mutual benefit shared by others, between species this is facilitation with service sharing or access to resources and within species, direct benefits by mutual benefits. These classifications provide a clear starting point for addressing the subject of helping behaviours.