Analysing body condition: mass, volume or density?

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1. Body condition (defined as the relative amount of energy reserves in the body) is an animal trait with strong ecological implications. In some animal taxa (e.g. arthropods), the external volume of the body part in which most nutrients are stored (e.g. abdomen) is used interchangeably with body mass to estimate body condition, making the implicit assumption that abdomen residual volume is a good surrogate of residual mass. However, the degree of correlation between these two measures should largely depend on the density of the nutrients stored. 2. We simulated two food-supplemented experimental groups of animals, each storing a slightly different amount of lipids either in their abdomens or in their entire bodies, and explored (i) how different estimates of condition were able to detect fixed differences between the groups; and (ii) how the amount of lipids stored could affect the outcome of non-intrusive measures of condition on a dichotomous variable (e.g. survival, mating success). We found that density body condition (body mass statistically controlled for structural body size and body volume) has much greater power to detect differences between experimental groups or effects on binary response variables than do classic mass/size or volume/size condition indices. 3. Using data on Lycosa tarantula (L.), a burrowing wolf spider, we report dramatic differences among these three indices in their ability to detect sex differences in the effect of feeding treatment on body condition at maturity. In particular, a plot of residual mass against residual volume reflecting nutrient density suggests that poorly fed spiders are nutritionally unbalanced, since well-fed spiders invest in nutrients of very different density. 4. Furthermore, using data on Scathophaga stercoraria (L.), the yellow dung fly, we found that an index of density condition was better at distinguishing condition differences among three populations than were mass or volume condition estimates alone. 5. We propose that including these three surrogates of condition (mass, volume and density) will substantially improve the accuracy of non-intrusive estimates of body condition, thus providing more powerful tools with direct application in a wide range of disciplines.