Measuring the shape of mortality across animals and plants: Alternatives to H entropy metrics reveal hidden type IV survivorship curves and associations with parental care at macro‐ecological scales

Abstract

The shape of mortality, or how mortality is spread across an organism's life course, is fundamental to a range of biological processes, with attempts to quantify it rooted in ecology, evolution, and demography. One approach to quantify the distribution of mortality over an organism's life is the use of entropy metrics whose values are interpreted within the classical framework of survivorship curves ranging from type I distributions, with mortality concentrated in late life stages, to type III survivorship curves associated with high early stage mortality. However, entropy metrics were originally developed using restricted taxonomic groups and the behavior of entropy metrics over larger scales of variation may make them unsuitable for wider‐ranging contemporary comparative studies. Here, we revisit the classic survivorship framework and, using a combination of simulations and comparative analysis of demography data spanning the animal and plant kingdoms, we show that commonly used entropy metrics cannot distinguish between the most extreme survivorship curves, which in turn can mask important macroecological patterns. We show how using H entropy masks a macroecological pattern of how parental care is associated with type I and type II species and for macroecological studies recommend the use of metrics, such as measures of area under the curve. Using frameworks and metrics that capture the full range of variation of survivorship curves will aid in our understanding of the links between the shape of mortality, population dynamics, and life history traits.