Population is sampled 2 or more times with previously unmarked individuals receiving a unique mark
'Captured' individuals are released back into the population
** Capture may be accomplished by physical capture, photography, DNA fingerprinting, etc.
Lincoln-Petersen method:
1 recapture event
Closed population
Equal capture probability of individuals during BOTH capture events
Marks are not lost, gained, or overlooked
K-sample Closed Population Models:
>1 (K) recapture/marking events
Closed population
Equal probability of capture within a given sampling occasion BUT probabilities can vary between sampling periods
Capture history for each individual is recorded
Program CAPTURE
Models to accommodate different capture probabilities:
Equal capture model (M0): Every animal has the same capture probability every time.
Heterogeneity model (Mh): Each animal (j) has a unique capture probability (pj)
'Trap' response model (Mb): All individuals have same initial capture probability (p) and all marked individuals in subsequent captures have a constant recapture probability (c). p and c can be different
Behavior heterogeneity model (Mbh): Each animal has its own unique pair of initial capture (pj) and recapture probabilities (cj)
Time variation (Schnabel) model (Mt): Every individual in the population has the same (re)capture probabilities that can vary over time (t). Thus, capture probabilities are (pt) for t = 1, 2, ..., K sampling occasions.
Time-heterogeneity model (Mth): Each individual has a unique capture probability that can vary through time (ptj)
Time-behavioral response model (Mtb): Every animal that has not been previously marked has an initial capture probability that can vary with time (pt) and every animal that has been previously caught has a recapture probability that can vary with time (ct)
Time-behavioral-heterogeneity model (Mtbh): Each individual has a unique initial capture probability that can vary with time (ptj) and each individual has a unique recapture probability that can vary with time (ctj).
** Each of these models have substantially different number of parameters that need to be estimated
** Use BOTH biological and statistical (e.g., AIC) arguments for model selection (additional notes on AIC)
Jolly-Seber method:
Jolly (1965) and Seber (1965) submitted idea separately
Every animal present in the population (marked and unmarked) has the same probability of capture during each sampling occasion (pt)
Every marked animal present in the population immediately after the tth sample has the same probability of survival (Øt) until the (t + 1)th sampling time.
Marks are not lost or overlooked.
Sampling time is negligible in relation to intervals between samples (i.e., demographic closure is satisfied during each sampling occasion) and each release is made immediately after the sample.
Every animal in the population is equally likely to emigrate and emigration is permanent
Model estimates population size, "mortality" rates (i.e., deaths + emmigration), and "births" (i.e., births + immigration)
Notation for the Jolly-Seber model:
Nt = total number of animals in the population at time (t) of the sample
Mt = total number of marked animals in the population at time (t) of the sample
mt = Number of marked animals caught in sample t
nt = Total number of animals caught in sample t
Rt = Total number of animals released after sample = nt - accidental deaths or removals
rt = Number of individuals released at sample t and caught again in some later sample
Zt = Number of individuals marked before sample t, not caught in sample t, but caught in some sample after sample t
To estimate population size: Program JOLLY (http://www.mbr-pwrc.usgs.gov/software)
Example From Krebs, C. J. 1989. Ecological Methodology.
Capture history for field voles: