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ROSS <br />Sexually mature white suckers in the Mill River (Amherst-Hadley, MA) spawned <br />less frequently than on an annual basis. Biased sex ratios favored males in the <br />spawning runs of this population; mature males were approximately three times <br />as likely to spawn in a given year as were females. The spawning frequency of <br />females increased with age, supporting the hypothesis that organisms exhibiting <br />reduced reproductive effort as part of their life history should increase repro- <br />ductive effort with increasing age. Excellent growth of white suckers in this <br />watershed indicates that their reproductive schedule probably is adaptive and is <br />not the outcome of poor nutritional or physiological status. <br />ITEROPAROUS fishes generally have been <br />thought to spawn annually. Exceptions to <br />this pattern have been documented for only a <br />few taxa: acipenserids (Roussou, 1957; Dads- <br />well, 1979); the paddlefish Polydon spalhula <br />('Meyer 1960); the striped bass Aforone saxalilis <br />(Jackson and Tiller, 1952); and several Eurasian <br />species of salmonids, cyprinids and silurids <br />(Maksunov, 1971). Here, we document a non- <br />annual spawning cycle for the white sucker Ca- <br />loslomus commersoni, and analyze the sex and age <br />specific characteristics of spawning frequency <br />in this species. <br />The white sucker is a widely distributed, long- <br />lived, ?North American freshwater species that <br />exhibits great variation in growth rate, age and <br />size at maturity, and fecundity (Carlander, 1969; <br />Beamish, 1973; Scott and Crossman, 1973). <br />White suckers are usually potadromous with <br />adults migrating up tributary streams to spawn <br />in early spring. Early studies suggested that the <br />sex ratio of fishes participating in a spawning <br />migration was 1:1 (Spoor, 1938; Raney and <br />Webster, 1942). However, biased sex ratios have <br />been measured in spawning migrations by other <br />investigators (Dente, 1948; Bouchard, 1955; <br />Geen et al., 1966). <br />Numerical dominance of one sex among <br />spawners could be caused by: 1) differential <br />mortality of the sexes, 2) deferred maturity of <br />one sex, or 3) a greater tendency for members <br />ofone sex to spawn on a non-annual basis. Olson <br />and Scidmore (1963), monitoring the later por- <br />tion of a spawning migration, found that suck- <br />ers fin-clipped two years previously outnum- <br />bered those marked during the previous <br />spawning run by nearly 2:1. Geen et al. (1966) <br />found that at least 22% of repeat spawners had <br />apparently skipped at least one spawning op- <br />portunity over a five-year period. The repeat <br />spawners were not analyzed by sex or age. These <br />studies indicate that suckers in some popula- <br />tions may reproduce on a non-annual basis. A <br />biased sex ratio among spawners could result in <br />such populations if one sex tended to spawn less <br />frequently than the other. However, none of <br />the above studies characterized the population <br />from which the spawners originated. <br />Reproductive effort that produces less than <br />the immediate, apparent-maximum number of <br />offspring could evolve if the number of surviv- <br />ing offspring produced in the lifetime of an in- <br />dividual is potentially enhanced (Williams, 1966; <br />Gadgil and Bossert, 1970; Goodman, 1974). Bull <br />and Shine (1979) have suggested that a corre- <br />lation exists between "low frequency reproduc- <br />tion" and "accessory reproductive activities" <br />(including breeding migrations, egg brooding <br />and live bearing) among iteroparous poikilo- <br />thermic vertebrate species. They postulated that <br />lifetime fecundity can be increased by non-an- <br />nual spawning if the associated reproductive ac- <br />tivities significantly increase the mortality rate <br />or require storage of energy which could be <br />otherwise channelled into gametes. <br />Reduced reproductive effort should be of <br />greatest benefit to younger age classes that could <br />increase fitness most by increasing the proba- <br />bility of producing offspring in future repro- <br />ductive seasons; thus, reproductive effort should <br />increase with age (Williams, 1966; Gadgil and <br />Bossert, 1970; Schaffer, 1974). Such an increase <br />in reproductive effort has been demonstrated <br />for a few fishes (Stearns, 1976). If white suckers <br />spawn on a non-annual basis and the reproduc- <br />tive schedule is adaptive, one might expect to <br />1985 by she American Society of Ichthyologists and 1lerpemlogists <br />Itl <br />I `u <br />r?,i?