McGUKK: PACIFIC HERRING OTOLITH KINGS 



the difference arose from factors that have already 

 been reported to affect the rate of deposition of nor- 

 mal rings. These factors include temperature 

 (Taubert and Coble 1977; Marshall and Parker 

 1982), short-term temperature fluctuations 

 (Brothers 1978), and feeding activity (Uchiyama and 

 Struhsaker 1981; Neilson and Geen 1982). Lough et 

 al. (1982) suggested that the first class of thin rings 

 were related to the inability of first-feeding larvae to 

 meet their metabolic energy demands during the 

 transition from yolk to exogenous food. This argu- 

 ment implies that the 1980 herring larvae were less 

 able to capture sufficient food during first feeding 

 than the 1981-82 larvae. However, this hypothesis 

 does not explain the presence of the faint rings in the 

 1980C larvae that were starved from hatch. 



Results of this study confirm the observations of 

 Geffen (1982) that the rate of normal ring production 

 is not always daily in young herring larvae and that it 

 is positively correlated with the rate of growth in 

 body size. The correlation means that normal rings 

 cannot be used with confidence to age wild herring 

 larvae less than about 20 mm long, unless the average 

 growth rate of the population is known to be higher 

 than about 0.36 mm/d (calculated from the regres- 

 sion of ring deposition rate on growth rate for Pacific 

 herring only). Growth of larval fishes is influenced by 

 several factors: temperature (Kramer and Zweifel 

 1970), food density (Haegele and Outram 1978), and 

 container size (Theilacker 1980). The tendency for 

 larger containers to support higher growth rates in 

 the four fed groups of this study may explain why only 

 two of the four had a daily ring pattern. The correla- 

 tion implies that, if the rate of growth is slowed or 

 stopped by starvation after a period of feeding, then 

 the rate of ring deposition should also slow or stop. 

 The two experimental groups that were treated in 

 this manner did not produce rings at rates that were 

 significantly different from those of their parent 

 feeding groups. This suggests that a starvation 

 period >5-8 d is necessary in order to demonstrate a 

 statistically significant effect. Larger rearing con- 

 tainers are also recommended to produce greater 

 contrast in growth rates between feeding and starv- 

 ing fishes. 



Container size, temperature, or prey size may 

 possibly have additional effects on the rate of ring 

 deposition apart from that which is explained by 

 growth rate. Temperature does explain some of the 

 residual variance of the ring deposition rate-growth 

 rate regression. However, published evidence on 

 effect of constant temperature on ring deposition 

 does not support the hypothesis that higher tem- 

 peratures produce more increments. For example, 



Neilson and Geen (1982) found no difference be- 

 tween the number of increments produced by 

 juvenile chinook salmon, Oncorhynchus tshawytscha, 

 reared at 5.2°C and at 11.0°C. The effects of such 

 environmental factors as light, temperature, and 

 prey type on the ring pattern of herring sagittae can 

 only be determined with a well-controlled ex- 

 perimental study. 



ACKNOWLEDGMENTS 



I gratefully acknowledge the assistance given by the 

 staff of the Bamfield Marine Station. I also thank 

 Gary Kingston for assistance in rearing fish in 1980, 

 Jeff Marliave for advice on rearing healthy marine 

 fish larvae, and Steve Campana for discussions on 

 grinding and reading fish otoliths. The paper has 

 benefited substantially from reviews by two 

 anonymous reviewers. This study was funded by a 

 G.R.E.A.T. Award from the Science Council of 

 British Columbia and by a grant to N.J. Wilimovsky 

 from the National Sciences and Engineering 

 Research Council of Canada. 



LITERATURE CITED 



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