Change-in-ratio methods for estimating population size

Change-in-ratio (CIR) methods can provide an effective, low cost approach for estimating the size of wildlife populations. They rely on being able to observe changes in proportions of population subclasses that result from the removal of a known number of individuals from the population. These methods were first introduced in the 1940’s to estimate the size of populations with 2 subclasses under the assumption of equal subclass encounter probabilities. Over the next 40 years, closed population CIR models were developed to consider additional subclasses and use additional sampling periods. Models with assumptions about how encounter probabilities vary over time, rather than between subclasses, also received some attention. Recently, all of these CIR models have been shown to be special cases of a more general model. Under the general model, information from additional samples can be used to test assumptions about the encounter probabilities and to provide estimates of subclass sizes under relaxations of these assumptions. These developments have greatly extended the applicability of the methods. CIR methods are attractive because they do not require the marking of individuals, and subclass proportions often can be estimated with relatively simple sampling procedures. However, CIR methods require a carefully monitored removal of individuals from the population, and the estimates will be of poor quality unless the removals induce substantial changes in subclass proportions. In this paper, we review the state of the art for closed population estimation with CIR methods. Our emphasis is on the assumptions of CIR methods and on identifying situations where these methods are likely to be effective. We also identify some important areas for future CIR research.