<br />4 Abundance Trends and the Status of the little Colorado River Population of Humpback Chub 1989-2006
<br />
<br />and Ryel, 1995). Details about these sampling programs are
<br />provided by Coggins and others (2006a).
<br />
<br />Tagging-Based Metrics
<br />
<br />The heart oftagging-based assessment is the large num-
<br />ber of uniquely tagged sub-adult [l50-199-rnm total length
<br />(TL)] and adult G::200-mm TL) fish that have been captured,
<br />measured. ,md implanted with pa5sive integrated transponder
<br />(PIT) tags. Since 1989, more than 19,000 HBC have been
<br />captured, tagged, and relea5ed with unique identitiers. These
<br />data are maintained in a central database housed at the USGS
<br />Grand Canyon Monitoring and Research Center.
<br />Mark-recapture methods to assess population abundance
<br />and vital rates have been widely used in fisheries and wildlife
<br />studies for more than 50 years, and numerous reviews have
<br />been conducted highlighting the general approaches (e.g.,
<br />Seber, 1982; Williams and others, 2001). Traditional methods
<br />(e.g., Jolly-Seber-type methods) generally rely on recaptures
<br />of tagged individuals to estimate abundance, recruitment, and
<br />survival. Basically, the approach is to create a known popula-
<br />tion of marked, or tagged, fish that are repeatedly sampled to
<br />obtain time series estimates of mark rate (i.e., the proportion
<br />of the overall population that is marked) and the number of
<br />marked fish alive in the population. These metrics are sub-
<br />sequently used to estimate capture probability, abund,mce,
<br />recruitment, and survival.
<br />The ASMR model ditfers from the traditional approach,
<br />because, in general, it contains more structural assumptions
<br />through the specification of a population accounting structure
<br />that governs transition of both marked and unmarked animals
<br />through ages and time. Age-structured stock assessment theory
<br />(Edwards and Megrey, 1989) is used to annually predict the
<br />numbers of marked and unmarked fish available for capture
<br />in a standard fisheries virtual population analysis framework
<br />(Quinn and Deriso, 1999). The total number of marked fish
<br />depends on the number of fi sh recently marked as well as the
<br />number of previously marked fish decremented by mortality
<br />rate. The number of unmarked fish depends on the recmitment
<br />over time, the number of fish marked from a given brood-year
<br />cohort, and the mortality rate. These annual predictions of the
<br />abundance of marked and unmarked fish are further segregated
<br />by age sucb that age-specific survival and capture probability
<br />may be modeled. Parameters are estimated by comparing pre-
<br />dicted and observed age- and time-specific captures of marked
<br />and unmarked fish in a Poisson likelihood framework.
<br />The ASMR model has three different parameteriza-
<br />tions (ASMR 1-3) that vary in how the temlinal abundance is
<br />estimated and how age- and time-specific capture probability
<br />is modeled. Both ASMR 1 and ASMR 2 assume that age- and
<br />time-specific captute probability can be modeled as th~prod-
<br />uet of an annual overall capture probability multiplied by age-
<br />specific vulnerability. This is similar to the common param-
<br />eterization of fishing mortality in assessment models under the
<br />"separability assumption" (Megrey, 1989) and diminishes the
<br />size of the parameter set since it is not necessary to separately
<br />
<br />estimate each age- and time-specific capture probability.
<br />These models further assume that vulnerability is asymptotic
<br />with age. As such, vulnerability is assumed to be unity for
<br />fish age-6 and older and estimated only for the younger fish.
<br />Finally, annual age-specific vulnerabilities are assumed to
<br />be equal among each sampling period. as described above.
<br />Implicit in this assumption is that within a sampling period,
<br />annual age-specitic capture probabilities differ only as a scalar
<br />value related to the annual overall capture probability.
<br />The primary difference between ASMR 1 and ASMR
<br />2 is how tbe tcrminal abundances are calculated. ASMR I
<br />estimates an overall terminal-year capture probability and
<br />calculates age-specific tcrminal abundances (both marked
<br />and unmarked) as the ratio of age-specific catch (both marked
<br />and unmarked fish) and age-specific capture probability (i.e.,
<br />product of the terminal-year capture probability and sam-
<br />pling period 4 age-specific vulnerability), In contrast, ASMR
<br />2 treats age-specific terminal abundances up to age-13 as
<br />individual parameters. Terminal abundances for subsequent
<br />ages are estimated by applying age-specific survivorship to the
<br />age-13 abundance. This difference in formulations decreases
<br />the parameter count for ASMR 1 relative to ASMR 2 at the
<br />expense of a5suming that the vulnerability schedule in the
<br />terminal year is identical to the rest of period 4.
<br />ASMR 3 is the most general model; it makes no assump-
<br />tion as to the age- or time-specific pattern in capture prob-
<br />ability. The conditional maximum likelihood estimates of
<br />age- and time-specific capture probability are used to predict
<br />the age- and time-specific catch of marked and unmarked fisb.
<br />Full detai Is of each of the models are provided by Coggins and
<br />others (2006b).
<br />In addition to the ASMR assessments, the time series of
<br />the annual spring abundance estimates in the LCR are updated.
<br />Abundance ofHBC in the LCR greater than or equal to 150
<br />mm TL was estimated during the early ] 990s and 2001-06,
<br />using closed population models. These models included
<br />the CAPTURE suite of models (Otis and others, 1978) ,md
<br />Chapman-modified, Lincoln-Petersen. length-stratified models
<br />(Seber, 1982). The recent estimators use data collected annu-
<br />ally during two sampling occasions in the spring. Full details
<br />of the sampling ,md estimation methods are provided by
<br />Douglas ,md Marsh (1996) and Coggins and others (2006a).
<br />Coggins and others (2006b) recommended exploring the
<br />use of individual capture histories within the ASMR frame-
<br />work to reduce confounding between capture probability and
<br />mortality. Though the updated ASMR models presented in
<br />this report do not yet incorporate individual capture histories,
<br />they clo model recaptured fish by annual-tagging cohort with
<br />the intent of reducing parameter confounding by increasing
<br />the number of observations available for parameter estimation.
<br />In the non-tag cohort, or pooled, version of ASMR described
<br />above and by Coggins and others (2006b), age- and time-spe-
<br />cific predictions of recaptured fish are not separated by year of
<br />tagging. As an example, assume that ASMR 3 predicts that 50
<br />marked age-6 HBC should be captured in 2002. These 50 fish
<br />could be comprised offish tagged as age-5 in 2001, age-4 in
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