Disentangling the effect of growth from development in size-related trait scaling relationships.

In plant ecology, the terms growth and development are often used interchangeably. Yet these constitute two distinct processes. Plant architectural traits (e.g. number of successive forks) can estimate development stages. Here, we show the importance of including the effect of development stages to better understand size-related trait scaling relationships (i.e. between height and stem diameter). We focused on one common savanna woody species (Senegalia nigrescens) from the Greater Kruger Area, South Africa. We sampled 406 individuals that experience different exposure to herbivory, from which we collected four traits: plant height, basal stem diameter, number of successive forks (proxy for development stage), and resprouting. We analysed trait relationships (using standardized major axis regression) between height and stem diameter, accounting for the effect of ontogeny, exposure to herbivory, and resprouting. The number of successive forks affects the scaling relationship between height and stem diameter, with the slope and strength of the relationship declining in more developed individuals. Herbivory exposure and resprouting do not affect the overall height-diameter relationship. However, when height and stem diameter were regressed separately against number of successive forks, herbivory exposure and resprouting had an effect. For example, resprouting individuals allocate more biomass to both primary and secondary growth than non-resprouting plants in more disturbed conditions. We stress the need to include traits related to ontogeny so as to disentangle the effect of biomass allocation to primary and secondary growth from that of development in plant functional relationships.