Trade-offs in interspecific comparisons in plant ecology and how plants overcome proposed constraints

Comparative studies on plant species have not distinguished between two inherently different applications of the idea of a trade-off. In the first case, the theoretical trade-off between two variables leads to a trend line, about which there is a degree of scatter. Any two species in the study are expected, in theory, to show a true trade-off, i.e. feature A must decrease for feature B to increase. This I call the ‘trend line’ type of plot. In the second case, the species are expected to fill a considerable part of the space defined by two axes and the theoretical trade-off leads to a boundary line which limits the extent of the cloud of points. In this case, many interspecific comparisons are not expected to show evidence of a trade-off, but those on the boundary line are expected to do so. This I call the ‘boundary line’ type of plot. In both types of plot, species within a given clade may show a true trade-off, while large numbers of unrelated species do not. First, I undertake to show, by means of examples, the reality of the distinction made above, and to demonstrate that statistically significant and ecologically important negative correlations between features A and B in the ‘trend line’ type of plot can be accompanied by huge variation from the trend line. Second, for plots of the ‘boundary line’ type, I undertake to show that authors have not always tested sufficiently rigorously the constraint lines concerning combinations of supposedly incompatible tolerances, and have not allowed for the absence of certain combinations of unfavourable conditions in the field and consequent lack of selection pressure for the evolution of species with a combination of extreme tolerances. I review two examples of each kind of constrained evolution. First, I take the worldwide leaf economics spectrum and the negative correlation between growth rate in high light and survival rate in deep shade. Second, I review the supposed incompatibilities between shade tolerance and drought tolerance, and between waterlogging tolerance and drought tolerance. I conclude that the term ‘trade-off’ is used too loosely. The common lack of true trade-offs has made possible the species richness of present-day vegetation. Also, small numbers of species have evolved combinations of tolerances that might seem improbable. Not enough research is being aimed at understanding the mechanistic basis of variation about trend lines, and the crossing of supposed boundary lines.