Demographic estimation methods for plants with unobservable life-states

Demographic estimation of vital parameters in plants with an unobservable dormant state is complicated, because time of death is not known. Conventional methods assume that death occurs at a particular time after a plant has last been seen aboveground but the consequences of assuming a particular duration of dormancy have never been tested. Capture-recapture methods do not make assumptions about time of death; however, problems with parameter estimability have not yet been resolved. To date, a critical comparative assessment of these methods is lacking. We analysed data from a 10 year study of Cleistes bifaria, a terrestrial orchid with frequent dormancy, and compared demographic estimates obtained by five varieties of the conventional methods, and two capture-recapture methods. All conventional methods produced spurious unity survival estimates for some years or for some states, and estimates of demographic rates sensitive to the time of death assumption. In contrast, capture-recapture methods are more parsimonious in terms of assumptions, are based on well founded theory and did not produce spurious estimates. In Cleistes, dormant episodes lasted for 1-4 years (mean 1.4, SD 0.74). The capture-recapture models estimated ramet survival rate at 0.86 (SEsimilar to0.01), ranging from 0.77-0.94 (SEsless than or equal to0.1) in any one year. The average fraction dormant was estimated at 30% (SE 1.5), ranging 16-47% (SEsless than or equal to5.1) in any one year. Multistate capture-recapture models showed that survival rates were positively related to precipitation in the current year, but transition rates were more strongly related to precipitation in the previous than in the current year, with more ramets going dormant following dry years. Not all capture-recapture models of interest have estimable parameters; for instance, without excavating plants in years when they do not appear aboveground, it is not possible to obtain independent time-specific survival estimates for dormant plants. We introduce rigorous computer algebra methods to identify the parameters that are estimable in principle. As life-states are a prominent feature in plant life cycles, multistate capture-recapture models are a natural framework for analysing population dynamics of plants with dormancy.