Modeling survival and mark loss in molting animals: recapture, dead recoveries, and exuvia recoveries

Capture-mark-recapture (CMR) analyses aim primarily at estimating relevant life history parameters, despite the fact that some individuals are not always recaptured, even if alive on the study site. Applying such approaches to species with a complex life cycle, such as insects, remains challenging because each change of stage tends to cause mark loss through molting. We developed a multistate model based on three exclusive events ("dead", "surviving and molting", and "surviving and staying in the same larval stage") to estimate probabilities of survival and mark loss. Estimates of biologically relevant parameters were derived from those of the probabilities of transition between these states. The model was applied to data from radio-tracking diodes glued on grasshoppers. The estimates of recapture probabilities decreased throughout the season for animals remaining alive, while the detection of dead animals and lost diodes was exhaustive. The survival probability was higher for larvae than for adults (0.98 vs. 0.96), and mark loss was stronger in larvae than in adults (0.09 vs. 0.06). We show that the survival rate of a species with a high rate of mark loss can be estimated using multistate models, provided that marks can be recovered after being lost. These models are flexible enough to test for several effects that potentially affect survival and mark loss probabilities.