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<br />22 <br /> <br />too small for individual tagging, and thus that <br />one may encounter problems separating <br />marked members from those that were marked <br />on previous occasions. If this is not a problem, <br />clipping of the adiposal fin may be the best <br />marking method. For short term studies, other <br />fins may be used, or various branding <br />methods. <br /> <br />The removal method <br /> <br />The second type of method used to estimate <br />population size in a small, closed area is the <br />removal method. This is the method most widely <br />used to estimate the population size of fish in <br />streams or the litoral zone of lakes when using <br />electrofishing and can be used if the capture is <br />efficient enough to reduce the population size <br />substantially. <br />To apply the removal method, the following <br />assumptions should be fulfilled (Seber, 1973; <br />p.312): <br /> <br />(1) The population is closed <br />(2) Equal catchability for all individuals <br />(3) Equal catch ability among the removals <br /> <br />The general case of k removals <br />To get estimates of population size from succes- <br />sive removals, one method is to use the maximum <br />likelihood estimator developed by Moran (1951) <br />and Zippin (1956). As there is no general explicite <br />solution, Zip pin (1956, 1958) provided graphs by <br />which the population size and the catchability can <br />be estimated in the case of 3, 4, 5 and 7 removals. <br />If k = the number of removals, c1 , c2. . . ck = the <br />catch in each consequtive removal, <br />T = (c, + c2 + ... + ck) = total catch, p = catch- <br />ability, and q ::::: 1 - p, a more flexible method is <br />to let a computer estimate q by iterative solution <br />of the expression <br /> <br /> k <br />^ kqk L (i - l)cj <br />q i = 1 (6) <br />= <br />^ 1 ^k T <br />P -q <br /> <br />In (6), the sum in the last term is C2 + 2c) for <br />k = 3, and C2 + 2c) + 3c4 for k = 4 and so on. If <br />(6) is used for iterative solution, a first guess of <br />p = c,/T can be used. The population size y is <br />estimated by using the q value obtained in the <br />expreSSIOn <br /> <br />^ T <br />Y = (1 - qk) <br /> <br />(7) <br /> <br />The sampling variances of y and pare <br /> <br />^ y(1- qk)qk <br />V(y) = (1- qk)2 _ (pk)2qk-' <br /> <br />(8) <br /> <br />and <br /> <br />r <br /> <br />v ^ _ (qP)2 (1 - qk) (9) <br />(p) - y [q(1 - qk? _ (kP)2qk] <br /> <br />The Standard Errors are the square roots of (8) <br />and (9). For confidence limits, see (3). <br />Although (6) and (7) are readily programmed <br />into a computer, it is sometimes convenient, e.g. <br />in the field, to use simpler methods. One such, <br />yielding practically the same result as the method <br />above, and which to our knowledge has not been <br />previously used, is the following. An estimator of <br />q is(l) <br /> <br />^ T - c1 <br />q= <br />T - ck <br /> <br />(10) <br /> <br />with the notation above. To obtain population <br />size, (7) is used. <br />Another simple method is Hayne's (1949) gra- <br />phical regression method. The relations (10) and <br />(7), however, are as simple to use, less subjective <br />and therefore recommended in favour of Hayne's <br />method. <br /> <br />.* <br />( <br /> <br />1 <br /> <br />, <br /> <br />The case of 3 removals <br />Junge and Libosvarsky (1965) found explicite <br />solutions of y and p in the case of k = 3. As this <br />case is of special interest, the solutions are given <br /> <br />t. <br />( <br /> <br />(I) Derived in appendix I. <br />