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<br />240 <br /> <br />COGGINS ET AL. <br /> <br />to represent a range of possibilities for explaining the <br />observed changes in catch rate in the LCR as well as <br />the ontogeny of downstream migration. This is <br />unsatisfactory as a long-term solution to the problem <br />of disentangling movement and sampling intensity <br />effects, but it allowed us to determine whether <br />recognition of seasonal movement results in different <br />abundance trend estimates than those obtained from the <br />spatially aggregated methods. The results of these <br />scenario tests were essentially the same as those of the <br />annual ASMR and Jolly-Seber models shown in Fig- <br />ure 5. We were not able to fit multistate models in <br />MARKJor reasons similar to those described above for <br />ASMR: low capture probabilities between states and <br />the associated failure to estimate a transition probabil- <br />ity. <br /> <br />Discussion <br />Population Trend and Abundance Assessments <br /> <br />All of our evidence implies that the adult humpback <br />chub population in the LCR has experienced a large <br />decline in abundance since 1989. Low sampling effort <br />in the late 19908 (1996-1999), heterogeneity in capture <br />probability related to age, and a large number of age- <br />classes make complicated models with large numbers <br />of parameters necessary to realistically estimate <br />population parameters. However, population trend <br />and annual population size estimates from all catch <br />indices and methods indicate declines in population <br />size of 30--60% since the early 1990s. <br />A comparison of the recent closed-population <br />(Lincoln-Petersen) estimates with other closed-popu- <br />lation estimates from the early 1990s indicates large <br />declines in population size. The only published <br />population estimates for humpback chub in the LCR <br />are closed-population estimates of fish exceeding 150 <br />nun TL by Douglas and Marsh (1996). Their estimates <br />for the spring of 1992 ranged from about 4,300 to <br />5,500 (Figure 3). Our closed-population estimates from <br />spring 2000-2003 ranged from about 2,000 to 3,400 <br />fish (TL > 150 nun; Figure 3). A comparison of <br />closed-population estimates for. fish occupying the <br />inflow reach suggests a decline in adult abundance of <br />about 60% between the early 1990s and 2001 (Valdez <br />and Rye11995; Trammell and Valdez 2003). Although <br />the assumptions of closed-population models (particu- <br />larly the lack of animals moving into and out of the <br />sampling area) may not have been met, Lin- <br />coln- Petersen estimates do allow some relaxation of <br />this closure assumption (pollock et al. 1990). For <br />example, if unmarked animals immigrate into the <br />sampling area, then the Lincoln-Petersen population <br />estimate is not biased for the second sample. If both <br />marked and unmarked animals emigrate between the <br /> <br />samples, then the Lincoln-Petersen estimate is un- <br />biased by emigration for the first sample. IT only <br />untagged animals emigrate between samples, then <br />population size estimates would be negatively biased. <br />However, it is unlikely that tagging effects only <br />occurred in the early 2000s, and any bias caused by <br />tagging probably occurred across all closed-population <br />estimators in the 1990s and 2000s. As such, these <br />estimates still serve as a relative indicator of an <br />apparently large decline in the humpback chub <br />population. <br />Using open-population models (Jolly-Seber and <br />ASMR), .the estimated LCR humpback chub popula- <br />tion of adult fish (i.e., TL >200 mm, age 4 and older) <br />is currently between 2,400 and 4,400 individuals <br />(Figure 5). No other open-population size estimates <br />have been published for this endangered species. The <br />TSM open-population trend estimates provide an <br />alternative to the age-dependent Jolly-Seber and <br />ASMR model annual population estimates. Population <br />growth estimates using TSM methods are robust to <br />heterogeneity in capture probability under the assump- <br />tion that the heterogeneity of the population does not <br />change over time (e.g., capture rates for smaller fish are <br />always lower than those for larger fish; Williams et al. <br />2002). Estimates of population growth also do not <br />depend on geographic closure of the sample area as <br />long as the measured population change of the sampled <br />area matches that of the population as a whole <br />(Schwarz 2001; Hines and Nichols 2002). All bi- <br />ologically reasonable TSM models, including con- <br />strained models that allow survival and capture <br />probability to remain time dependent and population <br />change to be time independent (Franklin 2001), <br />indicate annual declines in population size of up to <br />14%. IT the trends in annual estimated population size <br />from open-, closed-, or TSM population models are <br />examined, all approaches agree that the abundance of <br />the LCR humpback chub population has declined <br />significantly since at least the early 199Os. <br /> <br />Model Performance and Potential Assessment Errors <br /> <br />Abundance trends from the Jolly-age and ASMR <br />models are similar, though the absolute abundance <br />estimates differ between the two approaches, particu- <br />larly in the early years of the study. These differences <br />arise because the ASMR model predicts the number of <br />fish available for capture by using the existing age <br />structure of the population at the beginning of the study <br />while the Jolly-age approach only uses information <br />gained from recapture of tagged fish. Thus, as the <br />tagged-fish population increases in the middle to late <br />years of the study, the estimates from the two <br />approaches converge (Figure 4). <br />