34 in reduction of the sample size from 154 to 83 fish, providing the needed reduction in time needed for processing. Treat.ent of Data Modele for the relationship of fecundity to total length were obtained by least squares linear regression uaing the Minitab (TM) computer software package. The models utilized mean lengths and estimated fecundities for fish in 10 ma length intervals (170-179, 180-189, etc.), transformed to base 10 logarithms. Thi. procedure minimized bias due to unequal numbers in aome intervals and provided more uniform weighting of data over the total length range. Confidence intervals were calculated at the 95~ level for the slopes, intercepts, and predicted meana of the various .odel •• Age Determination Pla.tic impre •• ione of scalee were made uaing an Ann Arbor roller pr•••• Scale images were proJect.d by a tri-.implex micropro)ector with 40X magnification, and ages were determined u.ing standard method •• 35 RESULTS AND DISCUSSION Regression models were individually developed £or 1985 volumetric estimates, 1985 gravimetric estimates, 1986 volumetric estimates, and 1986 gravimetric estimates. Comparisons o£ the 95% con£idence intervals o£ the slopes and intercepts o£ the £our regression equations revealed overlap among all intercepts and slopes, respectively (Figures 3 and 4). The overlap was interpreted as evidence o£ no eigni£icant di££erences; t~ere were no signi£icant di££erences among the compared slopes or intercepts, allowing the conclusion that regression models were statistically similar. Since there were no signi£icant di££erences among the models within or between years, the 1985 and 1986 data were pooled £or each method o£ estimation, and a regression model was £ormulated £or each method. discussion o£ results re~ers From this point on, all to the combined 1985-86 data. The length-£requency distribution o£ the 83 £ish used in £ecundity analysis shows that the distribution is skewed toward the smaller sizes (170-229 mm) (Figure 5). This illustrates why means o£ each 10 mm length claas were used in £ecundity analysis to achieve more equal weighting o£ data over the length range £rom 170 to 299 mm (see Methods). A comparison was made between the mean lengths by age in the current study and the mean back-calculated lengths at annulus £or a larger sample o£ the perch population in 1985 and 1986 (Gallinat 1987) <Table 2). Since the £ish in the 36 86 G 85 -7.00 ~ GI -6.00 ---+-----1 -5'.00 -4.00 -3.00 INTERCEPT VALUE -2.00 -1.00 0.00 37 86 I G Iss 86 V I I I 185 I 2.00 2.50 3.00 G 3.50 SLOPE 4.00 V 4.50 5.00 38 ~ -.. 0'"• I I I I ... "J ~. t-r--......j .-. >I:.,) :z: lJJ =:. If) UJ ~ ~ (\J ----. I Qi I M O! i.L i (\J - -- -- ----- :z: (\J ....... UJ -~- - - ----- -- - -- - - .,......... ~I ------- - a:::: 1LJ ca (\J - - -- -- ---~-- tSt 00 ".0 V ':'-J is,I "!"'""'I .....t -- l!1 ...:.. W --J lSI 1-' r- I ~ :::I: I- :E ::. tSt 00 1".0 V Z ':\J "1"-1 "!"'""'I ,...... ---- .:.:- I --J .-<I:0 •r- 39 Table 2. Mean lengths by age £or the 83 £ish used in £ecundity analysis in the present study, compared to back-calculated lengths at annulus £or Gallinat's (1987) large sample collected during the same time period as the present study (1985-86). Length in mm (n in parentheses) Source Gallinat (1987) Present study III 154 (158) 187 (2) IV 192 (118) 199 (55) V 230 (44) 233 (23) VI 270 (6) 226 (2) 40 present study were collected early in the growing season, their lengths approximated the length at annulus. The mean lengths of ages IV+ and V+ fish in the current study agree closely with the corresponding back-calculated lengths at annulus found by Gallinat (1987). A similar comparison of ages lIl+ and VI+ is not meaningful because each class contained only two fish used for fecundity analysis. In addition, there was obvious selection for only large 111+ fish, since few fish were mature at this age (see Literature Review; Size at Maturity). The age-frequency distribution for the fish used in fecundity analysis reveals the vast maJority of the fish analyzed were age IV+ or V+ (Figure 6). Recall that Gallinat (1987) found in the same study area and time period as the present study, only were mature at age III, but lOO~ at age V (Figure 1). 27~ 86~ of the female yellow perch were mature at age IV and In addition, he showed that mortality was very high after age III. Thus, the first two year classes in which the maJority of the females were mature correspond to the predominant year classes of the fish analyzed in the present study. The small number of age 111+ fish in the sample is explained by the fact that very few feMales were mature at this age, and the saall number of age VI+ fish may readily be explained by high mortality of older £ish. The relatively small sample in the present study agrees closely with Gallinat's extensive data for the same population, implying that with regard to age, 41 >- + u z :::- L1J :=. c; w l..lJ 1-' Ct.: lJ... I ~:J: + ...... W I.!I :::- a:: Ct:: I.JJ m E I~ is) is) .:S) I~ I~ is) is) I~ I~ I~ =-Z'9""'l I~ "... I)) I'- ~.D uJ V 1"') .:\1 ~ 42 the fecundity models were based on a representative sample of the female spawning population. Regression Models The fecundity model derived from the volumetric estimates of fecundity (Appendix 5) is: log F where F' = = -4.0396 ~ fecundity and L (3.5834)10g L = total length. (r = 0.982)~ The correaponding fecundity model based on gravimetric estimstes of fecundity (Appendix 6) is: log F = -3.8258 ~ (3.5097)10g L (r = 0.986). Basic aimilarity of the models is apparent and both have high correlation coefficients. The models with the data points (means for 10 mm length intervals) used in their calcul&tion show the relationship graphically (Figure 7). Error Analysis It. is important to determine which model more accurat~ely describes the relationship of fecundity to length for the population. volumet~ric Evaluation of error analyais for and gravimetric estimates provides inSight to the accuracy of the models (Table 3). fish used in error analys~s. For each of the three the fecundity estimates were compared to the actual fecundity determined by the total egg count £or the fish. It~ is apparent that the volumetric method tended to underestimate slightly (-3.4~), tended to overestimate (+8.8~) while the gravimetric method (Table 3). Although the 43 44 Table 3. Comparison o£ £ecundity estimates to total egg counts. Deviations o£ the individual estimates £rom the actual counts appear in parentheses. Fecundity Fish 1.0. NUMber Length (mm) Actual Count Volumetric Estimate 18 988 (+5.6%) 25 430 (-5.1%) 27 197 (-10.6%) 20 860 (+16.0%) 28 685 (+7.0%) 31 414 (+3.3%) -3.4% (8.2%) +8.8% (6.5%) 85-39 202 17 977 85-17 227 26 808 85-2 242 30 425 Mean Error (Standard Deviation) Gravimetric Estimate 45 gravimetric method was slightly less variable <SD the volumetric method (SD 8.2~). 6.5~) than the volumetric estimatea were closer to the actual known number o£ eggs. The regression model based on volumetric .stimates thus appears to more closely describe the actual population than the gravimetric model. The volumetric model was therefore utilized exclusively in all additional analyses presented. Predictions From the Models Fecundity models may be used to predict the number o£ eggs produced by a £ish o£ a given length. In addition. i£ the mean length at a given age is known, the mean fecundity for that age class can be predic~ .d. This information may in turn be used in estimates o£ population fecundity (Bagenal 1978). Predicted fecundities and 95~ confidence intervals for yellow perch within the length range o£ the present study were calculated from the volumetric model (Table 4). Equivalent predictions were also made for each age class in the present study (Table 5). The mean length for each age class was calculated from Gallinat's (1987) data for 1985-86. The prediction for age 111+ is probably not meaningful. since the average fish in this age class would not be mature, as discussed earlier (see Figure 1). Comparison to Previous Studies The fecundity o£ Lake Michigan yellow perch has previously been reported by Brazo et ale (1975) for fish 46 Tabla 4. Fecundity predictions and 95% con£idence intervals £or yellow perch o£ the given lengths, calculated £rom the volumetric £ecundity model, log F = -4.0396 ~ 3.5834 log L. Fecundity Length (mil) 180 190 200 210 220 230 240 250 260 270 280 290 Prediction 11 13 16 19 22 26 30 35 41 47 53 60 010 370 070 140 610 510 880 740 140 090 650 840 c. 95% 9 11 14 17 20 24 28 32 36 41 46 52 630 950 620 620 940 520 360 460 860 580 660 100 - 12 14 17 20 24 28 33 39 45 53 61 71 I. 600 960 660 780 410 660 620 360 910 330 690 060 47 Table 5. Age 111+ IV+ V. VI· Fecundity predictions and 95% confidence intervals for yellow perch of the given ages. calculated from the volumetric fecundity model: log F = -4.0396 • 3.5834 log L. Mean Length 1 (mm) 154 192 230 270 Fecundity Prediction 6 13 26 47 300 880 510 100 95" C. I. 5 12 24 41 130 450 520 580 7 730 - 15 470 - 28 660 - 53 330 1 The mean length for each age class was calculated from Gallinat's (1987) data for 1985-86. 48 190-354 mm total length collected near Ludington, Michigan in 1972, and by Wells and Jorgenson (1983) £or 174-355 mm £ish collected at Saugatuck, Michigan in 1972 and 1979. The regression models o£ £ecundity versus total length obtained in these studies are: Brazo et al. = log F (1975): -3.712 + (3.451)10g L (no r value given) Wells and Jorgenson (1983): = log F -3.99 (3.56)10g L + (r = 0.96). Recall that the model best describing the population in the present study (i. e., the volumetric model) is log F = -4.0396 + (3.5834)10g L (r ~ 0.982). When the previously reported models are plotted on the same axes as the present model based on volumetric estimates (con£ined to the length rsnge o£ the present study) they £all within the (Figure 8). 95~ con£idence contour o£ the present model In £act, the model o£ Wells and Jorgenson is virtually identical to and superimposed on the current .odel. This indicates that over the length range compared, the £ecundity-length relation determined in the present study does not signi£icantly di££er £rom those reported previously £or Lake Mich1gan yellow perch by Brazo et al. (1975) and Wells snd Jorgenson (1983). It should be noted that the earlier studies (and the populations in general) included larger £ish than the present study. Although 1t has been stressed that it is i.portent to limit comparisons o£ regression equations to those generated £rom £ish o£ the same size, this caution 49 ~lOO. 000 BRAZO E!!'t al. ( .1 9"7 5 80.000 ) _._.- ( .1 9 8 3 ) _.-.- PRESENT STUDY PRESENT STUDY , I I I I 60.000 I I I I I I I I I I I I I I I 40.000 ~ I I !-< H I I I I 0 I I I I I I I I Z J U I I UJ I I II. I I 20.000 I I I , I I I I I ., I I I I I I I , I I I I , .10.000 I , I I 8. 000 a '.a .-4 a a N a ~ N TOTAL a a a (') 11l .-4 al N N LENGTH a a N a a a N (') ~ (MM) al N N 50 seems to be relevant only to claims of significant differences. The fact that over the length range of the present study, the previously reported regression equations fall within the 95~ confidence contour of the present model, is, i£ anything, even stronger support for a lack of significant differences than if comparisons were made among equations generated from fish of the same size. In addition to Brazo et ala (1975) and Wells and Jorgenson (1983>. regression models of fecundity versus length of yellow perch have been reported by Muncy (1962), Sheri and Power (1969), Tsai and Gibson (1971>, Sztramko and Teleki (1977). and Hartman et ala geographic locations. (1980), for various Unfortunately, the fecundity predictions of the present study are only directly comparable to those of Muncy (1962; Severn River, Maryland) and Hartman et ala (1980; Lake Erie), because the regressions in the other studies utilized fork lengths instead of total lengths. 1 Predicted fecundities over the length range of the present study were calculated from the regression models of Muncy (1962) and Hartman et ala (1980) (Table 6). values may be compared to the equivalent 95~ intervals for the present study (Table 4). These confidence It is apparent that while there are no Significant differences between the Lake Erie study of Hartman et ala (1980> and the present study, in Muncy's Severn River study the predicted fecundity 1. No standard conversion factor for yellow perch fork length to total length was found. 51 Table 6. Predicted yellow perch £ecundities over the length range o£ the present study, calculated £rom the regression models o£ Muncy (1962) and Hartman et al. (1980). Fecundity ---------------------------Total Length (mm) Muncy (1962) Hartman at al. (1980) --------------------------------------------180 190 200 210 220 230 240 250 260 270 280 290 5 6 8 10 12 14 16 20 23 27 31 36 500 800 400 300 400 800 900 000 400 300 700 500 11 14 17 20 24 28 33 38 44 50 57 65 600 100 000 300 100 300 000 300 200 600 800 600 52 is below the lower confidence limit of the present study over the entire length range compared. It is not clear. however. whether this represents a true difference. because Muncy's data include some fecundity estimates from eggs which were stripped from live fish. possibly resulting in inaccurately low predicted fecundities. The discovery that the fecundity of a given size yellow perch has not significantly changed in southern Lake Michigan since 1972 is somewhat surprising. As noted earlier. growth rates and length-weight relationships have declined dramatically in Indiana waters in recent years. apparently due to the current high population density (Gallinat 1987). Since fecundity is generally conSidered to be highly responsive to density-dependent factors and related influences such as food supply (see Literature Review). it was expected that fecundity would have decreased with increasing fish denSity. The apparent contradiction between decreased growth rates and constant fecundity suggests that some sort of threshold of denSity and/or food supply conceivably could be involved in influencing fecundity of yellow perch in southern Lake Michigan. this were the case. a moderate decrease in foo~ ~ncrease If in density and/or supply m1ght cause reduced growth rates without significantly affecting fecundity. Assuming that density and/or food supply in the study area have not crossed the hypothetical critical threshold. the hypothesis Just proposed would explain why fecundity of yellow perch in southern Lake Michigan has not significantly changed since 53 1972 despite decreased growth rates. Although the fecundity of a given size fish has apparently not changed with the increased density and reduced growth of yellow perch in the sample area, it is possible that the population fecundity has been altered due to a shift in the length-frequency distribution toward smaller fish. Subsequent research may provide insight ss to whether or not this has indeed occurred. 54 SUMMARY AND CONCLUSIONS Volumetric and gravimetric methods o£ £ecundity determination were conducted on 83 yellow perch (ranging in total length £rom 172 to 290 mm) collected £rom Lake Michigan near Michigan City, Indiana in 1985 and 1986. Major £indings o£ the project are as £ollows: 1. Linear regression models for £ecundity versus total length developed separately for 1985 and 1986 samples were not signi£icantly dif£erent from each other, based on overlap of the 95~ confidence intervals o£ the slopes and intercepts of the equations. The 1985 and 1986 data were there£ore pooled for each method o£ analysis. 2. The model for fecundity versus total length derived from the volumetric fecundity estimates was: (3.5834)10gL (r = 0.982). log F 3. + -4.0396 + The corresponding model derived from the gravimetric fecundity estimates was: -3.8258 = log F = (3.5097)10gL (r = 0.986). The volumetric estimates were determined to be more accurate than the gravimetric estimates by comparison to total egg counts o£ three ovaries. The volumetric model was therefore used for all subsequent analyses. 4. Predictions were made £or the £ecundities o£ yellow perch at 10 mm length intervals over the length range of the present study, utilizing the volumetric model. Mean lengths of the age classes represented were used to predict the mean fecundity o£ each age class. 55 5. The £ecundity o£ a yellow perch o£ a given length £rom 172-290 mm has not changed signi£icantly in southern Lake Michigan since 1972, despite the £act that population density has increased and growth rates have decreased. This was determined by comparing previously reported regression equations with the 95% con£idence contour o£ the present equation. 56 LITERATURE CITED Bagena1, T. B. 1978. Aspects of fiah fecundity. Pages 7~)-101 in S. D. Gerking, editor. Ecology of freshwater fish production. John Wiley and Sons, New York. Bagena1, T. B., and E. Braum. 1978. Eggs and early life h:lstory. Pages 165-201 in T. B. Bagenal, editor. Methods for assessment of fish production in fresh waters. Blackwell Scientific, Oxford. Becker. G. C. 1983. Fishes of Wisconsin, first edition. University of Wisconsin Press, Madison. Brazo, D. C., P. I. Tack, and C. R. Liston. 1975. Age, growth, and fecundity of yellow perch, Perca flavescens (Mitchill), in Lake Michigan near Ludington, Michigan. Transactions of the American Flsheries Society 104:726-730. Collette, B. B., and seven co-authors. 1977. Biology of the percids. Journal of the Fisheries Research Board o£ Canada 34:1890-1899. Dodgshun, T. J. fish eggs. 1980. Simple, accurate method of counting Progressive Fish-Culturist 42:237-238. Eddy, S., and T. Surber. 1960. Northern fishes, with special reference to the Upper Mississippi Valley, revised edition. C. T. Branford, Newton Centre, Massachusetts. El-Zarka, S. E. 1959. Fluctuations in the population of yellow perch in Saginaw Bay, Lake Huron. U. S. Fish and Wildlife Service Fisheries Bulletin 59:365-415. Emery, A. R. 1973. Preliminary comparisons of day and night habits of freshwater fish in Ontario lakes. Jc~urnal of the Fisheries Research Board of Canada 30: 761-774. Gallinat, M. P. 1987. Population analysis and food habits o£ the yellow perch, Perca flavescens (Mitchill), in Indiana waters o£ Lake Michigan, 1984-86. Master's thesis. Ball State University, MunCie, Indiana. Hartman, W. L., S. J. Nepszy, and R. L. Scholl. 1980. Minimum size limits for yellow perch <Perca flav.scens) in western Lake Erie. Great Lakes Fishery Commission Technical Report 39. 57 Healey. M. C. 1978. Fecundity changes in exploited populations of lake whitefish (Coregonus clupeaformis) and lake trout (Salvelinus namaycuah). Journal of the Fisheries Reaearch Board of Canada 35:945-950. Hokanson, K. E. F. 1977. Temperature requirements of some percids and adaptations to the seasonal temperature cycle. Journal of the Fisheries Research Board of Canada 34:1524-1550. Jobea, F. W. 1952. Age, growth, and production of yellow perch in Lake Erie. U. S. Fiah and Wildlife Service Fisheries Bulletin 52:205-266. Kucera, P. A., and J. L. Kennedy. 1977. Evaluation of a sphere volume method for estimating fish fecundity. Progressive Fish-Culturist 39:115-117. McComish, T. S. 1981. Yellow perch population characteri.tics in Indiana waters of Lake Michigan, 1976-79. Final Report of Federal Aid Project 3-283-R, Segment 3. McKeag, K. J. 1987. An evaluation of the potential forage fish populations in Indiana waters of Lake Michigan: 1973 vs. 1984-86. Master's thesis. Ball State University, Muncie, Indiana. Muncy, R. J. 1962. Life history of the yellow perch, Perca flavescens, in estuarine waters of the Severn River, a tributary of Chesapeake Bay, Maryland. Chesapeake Science 3:143-159. Newton, S. H., and R. V. Kilambi. 1973. Fecundity of the white ba •• , Morone chrysops (Rafinesque>, in Beaver Reservoir, Arkansas. Transactions of the American Fisheries Society 102:446-448. Ney, J. J. 1978. A synoptic review of yellow perch and walleye biology. Pages 1-12 in R. L. Kendall, editor. Selected coolwater fishes of North America. American Fisheries Society (Special Publication 11). Washington, D. C. Nielsen, L. A., and D. L. Johnson, editors. 1983. Fisheries techn~ques. American Fisheries SOCiety, Bethesda, Maryland. Nikolskii, G. V. 1969. Theory of fish population dynamics as the biological background for rational exploitation and management of fishery resources (translated from Russian by J. E. S. Bradley>. Oliver and Boyd, London. 58 Scott, W. B., and E. J. Crossman. 1973. Freshwater £ishes o£ Canada. Fisheries Research Board o£ Canada (Bulletin 184), Ottawa. Sheri, A. N., and G. Power. 1969. Fecundity o£ the yellow perch, Perca £lavescens Mitchill, in the Bay o£ Quinte, Lake Ontario. Canadian Journal of Zoology 47: 55-58. Sztramko, L., and G. C. Teleki. 1977. Annual variations in the fecundity of yellow perch from Long Point Bay. Lake Erie. Transactions of the American Fisheries Society 106:578-582. Thorpe, J. E. 1977. Morphology, physiology, behavior, and ecology of Perca fluviatilis L. and ~. flavescens Mitchill. Journal of the Fisheries Research Board of Canada 34:1504-1514. Treasurer, J. W. 1981. Some aspects of the reproductive biology of perch Perea fluviatilis L. fecundity, maturation, and spawning behavior. Journal of Fish Biology 18:729-740. Tsai, C., and G. R. Gibson, Jr. 1971. Fecundity o£ the yellow perch, Perca flavescens (Mitch!ll), in the Patuxent River. Maryland. Chesapeake Science 12: 270-274. Volodin, V. M. 1979. The fecundity of the perch, Perca fluviatilis, £rom Rybinsk Reservoir. Journal o£ Ichthyology 19:85-92. Wells. L. 1977. Changes in yellow perch <Perca £lavescens) populations of Lake Michigan, 1954-75. Journal of the Fisheries Research Board of Canada 34: 1821-1829. Wells, L., and S. C. Jorgenson. 1983. Population biology of yellow perch in southern Lake Michigan, 1971-79. U. S. Fish and Wildli£e Service Technical Paper 109. Wolfert, D. R. 1969. Maturity and fecundity of walleyes from the eastern and western basins of Lake Erie. Journal of the Fisheries Research Board of Canada 26: 1877-1888. 59 APPENDIX SECTION 60 Appendlx 1. loD. Number 85-1 85-2 85-3 85-4 85-5 85-6 85-7 85-8 85-9 85-10 85-11 85-12 85-13 85-14 85-15 85-16 85-17 85-18 85-19 85-20 85-21 85-22 85-23 85-24 85-25 85-26 85-27 85-28 85-29 85-30 85-31 85-32 85-33 85-34 85-35 85-36 85-37 85-38 85-39 85-40 85-41 85-42 Maximum total length, body weight, £resh ovary weight, and £resh ovary volume o£ £emale yellow perch collected in Indiana waters o£ Lake Michigan on May 8, 1985. Ages are also given £or selected £ish. Length (mm) 190 242 205 198 193 205 178 245 222 203 188 208 195 200 290 220 227 187 260 216 214 186 194 197 204 184 178 198 184 201 202 189 208 190 193 200 181 201 202 204 200 206 Body Weight Age (g) Ovary Weight <g) IV+ V+ 80 170 96 94 83 100 68 184 116 100 74 88 82 96 334 126 146 72 244 124 110 68 86 86 96 74 66 86 72 92 98 74 110 70 83 90 74 92 96 98 90 100 13.3 32.2 5.8 10.0 10.5 8.7 10.7 33.9 9.5 14.1 4.3 7.6 12.7 16.2 75.8 20.0 24.3 9.0 48.5 22.5 7.9 6.8 13.6 13.4 11.8 9.3 8.8 10.0 12.2 11.2 13.7 10.3 16.2 11.5 8.1 8.7 9.2 11.5 15.6 10.4 9.7 7.6 IV+ IV+ IV+ 111+ V+ IV+ 111+ V+ V+ V+ IV+ V+ IV+ IV+ IV+ IV+ IV+ IV+ IV+ IV+ V+ V+ IV+ IV+ IV+ Ovary Volume (mL) 13.0 35.0 5.5 10.0 10.0 8.5 11.0 33.0 9.0 14.0 4.0 7.0 12.5 16.0 73.0 19.0 23.0 8.5 47.0 22.0 7.5 6.5 13.5 13.0 11.5 9.0 9.0 9.5 12.0 11.0 13.0 10.0 15.5 11.0 8.0 8.0 9.0 11.5 15.0 10.0 9.5 7.0 ------~----------------------------------------------- ------ 61 Appendix 1 continued. 1.D. Number 85-43 85-44 Length (mm) 196 206 Body Weight Age (g) 76 84 Ovary Weight (g) 7.2 7.8 Ovary Volume (mL) 7.0 7.0 62 Appendix 2. Date of capture. maximum total length. body weight. and fresh ovary weight o£ £emale yellow perch collected in Indiana waters of Lake Michigan in 1986. Ages are also given £or selected fish. ----------------------------------------------------------Body Ovary 1.0. Number Date (mo./day> Length (mm) Age Weight (g) Weight <g) ----------------------------------------------------------86-1 86-2 86-3 86-4 86-5 86-6 86-7 86-8 86-9 86-10 86-11 86-12 86-13 86-14 86-15 86-16 86-17 86-18 86-19 86-20 86-21 86-22 86-23 86-24 86-25 86-26 86-27 86-28 86-29 86-30 86-31 86-32 86-33 86-34 86-35 86-36 86-37 86-38 86-39 86-40 86-41 86-42 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/8 181 203 176 220 203 190 202 205 228 197 180 218 188 229 172 207 204 192 216 211 211 211 212 193 183 180 250 199 202 196 198 202 209 240 181 197 191 208 185 176 190 174 IV+ IV+ IV+ IV+ V+ V+ IV+ IV+ V+ IV+ IV+ IV+ VI+ IV+ IV+ V+ IV+ IV+ V+ VI+ IV+ IV+ IV+ IV+ 65 86 71 120 92 79 97 100 142 88 64 122 79 102 60 104 95 80 108 95 101 113 106 83 69 67 194 86 99 91 81 93 98 151 70 91 74 107 69 69 80 66 9.6 15.0 12.8 17.6 14.3 11.8 19.0 15.5 18.8 13.3 10.6 21.4 15.6 15.3 8.8 22.4 15.5 13.0 20.5 14.9 12.1 22.9 14.5 14.5 12.0 12.5 38.7 13.5 13.4 14.6 13.3 14.9 12.5 25.1 12.5 15.1 12.6 19.1 11.9 11.0 13.2 9.9 ----------------------------------------------------------- 63 Append.lx 2 continued. ------~----------------------------------------------------- 1. D. Number Date (mo./dey) Length (mm) Age Body Weight (g) Ovary Weight <g) -----_._---------------------------------------------------86-43 86-44 86-45 86-46 86-47 86-48 86-49 86-50 86-51 86-52 86-53 86-54 86-55 86-56 86-57 86-58 86-59 86-60 86-61 86-62 86-63 86-64 86-65 86-66 86-67 86-68 86-69 86-70 86-71 86-72 86-73 86-74 86-75 86-76 86-77 86-78 86-79 86-80 86-81 86-82 86-83 86-84 86-85 86-86 86-87 86-88 5/8 5/8 5/8 5/8 5/8 5/8 5/8 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 191 192 186 209 205 180 175 173 189 242 239 184 245 283 301 244 240 223 242 182 250 208 225 185 186 224 202 201. 201 202 205 223 205 198 205 185 187 199 180 207 207 189 187 186 200 210 IV+ IV+ IV+ IV+ V+ V+ V+ V+ IV+ IV+ V+ IV+ V+ V+ IV+ IV+ IV+ V+ IV+ 77 85 77 99 103 70 55 52 65 202 163 67 191 254 363 191 176 139 151 60 198 103 124 73 75 143 84 96 109 101 104 134 106 93 101 78 81 100 69 105 109 80 76 74 99 112 6.8 12.8 13.0 14.3 17.1 10.7 7.3 10.7 13.8 55.3 31.4 11.7 32.2 40.5 89.7 32.8 29.5 28.0 21.4 9.6 41.4 18.4 22.7 9.9 16.1 26.3 19.0 21.0 22.8 18.2 18.3 20.8 16.8 18.2 18.6 14.4 18.7 16.6 10.4 19.8 16.5 14.1 11.2 14.5 20.0 19.8 ----------------------------------------------------------- 64 Appendix 2 continued. 1.0. Number 86-89 86-90 86-91 86-92 86-93 86-94 86-95 86-96 86-97 86-98 86-99 86-100 86-101 86-102 86-103 86-104 86-105 86-106 86-107 86-108 86-109 86-110 Date (mo./day) 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 5/27 6/4 Length (mm) 219 228 209 208 174 202 194 191 200 189 172 211 216 189 253 192 193 187 191 182 205 255 Body Weight Age (g) IV+ IV+ IV+ IV+ IV+ IV+ 108 161 107 100 62 92 93 88 89 87 63 106 124 81 221 76 76 71 84 72 101 219 IV+ IV+ IV+ V+ IV+ V+ Ovary Weight (g) 17.8 31.3 21.7 15.7 10.3 16.7 16.0 14.8 11.4 14.7 9.3 20.9 21.0 12.9 50.2 7.6 10.3 10.7 12.5 11.4 19.1 43.0 ------------------------------------------------------ ----- &5 Append~x 3. Data used in volumetric fecundity estimation. Egg Counts of Subsamples Fish I.D. Number Total Ovary Volume (mL) Sample a Sample b Sample c 85-1 85-2 85-4 85-5 85-6 85-7 85-8 85-9 85-1:3 4.65 9.95 3.53 3.81 2.95 3.62 10.80 4.40 4.00 22.17 6.50 7.15 2.33 16.52 7.29 1.93 4.50 3.72 3.24 2.95 3.81 3.38 5.15 2.64 2.90 4.70 3.72 4.30 5.55 4.30 4.35 7.29 7.24 5.55 2.85 4.55 7.44 5.70 3.91 8.18 4.65 4.00 14.76 4.65 5.40 4.90 317 265 384 393 462 308 313 535 352 255 396 344 338 309 346 499 359 464 424 327 331 324 457 549 383 399 464 343 330 440 509 372 265 376 303 254 289 361 300 385 310 298 264 265 293 376 323 274 435 415 516 313 335 529 371 286 408 365 362 316 338 513 352 474 433 315 315 330 458 576 398 418 460 340 333 435 518 381 273 378 305 258 276 355 310 389 304 301 261 266 303 401 333 281 421 435 483 321 325 549 384 274 417 358 363 325 346 503 373 451 420 346 330 329 477 592 404 395 481 342 341 449 522 385 268 371 315 258 283 350 299 394 321 298 274 268 286 390 85-1~5 85-11; 85-1'7 85-1,B 85-1'9 85-20 85-2:2 85-2'4 85-2'5 85-215 85-2'7 85-2'9 85-3.2 85-33 85-3'5 85-37 85-39 85-40 86-3 86-4 86-5 86-6 86-9 86-12 86-14 86-15 86-18 86-19 86-20 86-21 86-22 86-23 86-26 86-27 86-28 86-30 86-31 66 Appendix 3 continued. Egg Counts o£ Subsamples Fish 1 .. 0. Number Total Ovary Volume (mL) Sample a Sample b Sample c 86-3~l 8.08 6.90 4.00 3.38 3.24 4.40 3.53 2.03 2.44 4.40 20.33 10.85 12.90 12.70 10.90 9.37 7.34 2.90 16.06 7.24 9.27 7.93 3.62 6.20 6.30 5.75 10.24 6.90 5.10 3.24 4.65 3.24 5.25 7.73 15.85 3.67 14.36 322 266 343 314 356 282 335 404 347 379 166 235 242 245 209 257 235 260 254 190 231 289 329 330 254 344 222 288 391 279 293 396 250 328 253 310 229 347 272 335 315 349 277 332 401 358 370 175 252 235 254 195 252 225 253 256 205 216 310 320 341 245 363 220 303 394 246 296 414 256 320 260 306 219 333 292 337 319 361 303 326 387 348 375 169 235 246 258 203 270 232 253 258 201 242 315 322 341 247 344 220 302 395 268 296 403 255 330 241 297 220 86-3~~ 86-3~~ 86-4() 86-42 86-46 86-48 86-49 86-50 86-51 86-5:2 86-5~3 86-5~) 86-58 86-5~3 86-60 86-6:L 86-6:2 86-6:3 86-6~) 86-6l~ 86-7·~ 86-8~) 86-87 86-8l~ 86-8~3 86-90 86-9:L 86-92 86-9:3 86-9·~ 86-9<3 86-100 86-101 86-103 86-10? 86-1:LO 67 Appendix 4. Data used in gravimetric fecundity estimation. The egg counts of the subsamplea were the same for each fish as those given in Appendix 3. Dry Weights of Subsamples <g) Fish I.D. Number 85-1 85-2 85-4 85-5 85-6 85-7 85-8 85-9 85-13 85-15 85-16 85-17 85-18 85-19 85-20 85-22 85-24 85-25 85-26 85-27 85-29 85-32 85-33 85-35 85-37 85-39 85-40 86-3 86-4 86-5 86-6 86-9 86-12 86-14 86-15 86-18 86-19 86-20 86-21 86-22 86-23 86-26 86-27 Total Dry Ovary Weight (g) 1.5627 3.6013 1.0755 1.3500 0.9228 1.2547 3.6217 1.2824 1.3870 7.0287 2.0181 2.5811 0.7492 5.2615 2.3571 0.6571 1.5426 1.2669 1.0611 1.0342 1.3344 1.0915 1.7391 0.8685 1.0158 1.5159 1.2514 1.4738 1. 6799 1.4504 1.3959 2.4125 2.2088 1.9906 0.~680 1.3702 2.1532 1.7216 1.3898 2.7083 1.5556 1.2851 4.7444 Sample a Sample b Sample c 0.0295 0.0305 0.0266 0.0296 0.0282 0.0316 0.0300 0.0282 0.0295 0.0284 0.0282 0.0312 0.0285 0.0298 0.0306 0.0300 0.0309 0.0322 0.0298 0.0315 0.0320 0.0302 0.0306 0.0297 0.0313 0.0297 0.0298 0.0320 0.0281 0.0312 0.0290 0.0293 0.0283 0.0329 0.0301 0.0274 0.0269 0.0304 0.0319 0.0307 0.0299 0.0288 0.0307 0.0307 0.0317 0.0286 0.0300 0.0295 0.0304 0.0315 0.0274 0.0298 0.0300 0.0284 0.0327 0.0278 0.0304 0.0295 0.0317 0.0310 0.0319 0.0298 0.0301 0.0298 0.0301 0.0307 0.0306 0.0320 0.0302 0.0295 0.0316 0.0281 0.0314 0.0288 0.0297 0.0290 0.0329 0.0301 0.0285 0.0269 0.0297 0.0323 0.0306 0.0308 0.0286 0.0304 0.0312 0.0318 0.0283 0.0305 0.0281 0.0307 0.0309 0.0281 0.0312 0.0294 0.0285 0.0321 0.0283 0.0314 0.0299 0.0310 0.0324 0.0308 0.0290 0.0324 0.0281 0.0302 0.0312 0.0306 0.0327 0.0282 0.0309 0.0326 0.0291 0.0317 0.0285 0.0303 0.0288 0.0317 0.0308 0.0285 0.0272 0.0294 0.0316 0.0316 0.0317 0.0283 0.0318 ------------------------------------------------------ ----- 68 Appendix 4 continued. Dry Weights of Subsamples <g) Fish 1.0. Number 86-2~~ 86-3D 86-3:L 86-34 86-3~3 86-4() 86-42 86-4~~ 86-4'3 86-50 86-51 86-5:2 86-5:3 86-5~5 86-5j~ 86-5'3 86-60 86-6.1 86-6:2 86-6:3 86-6~5 86-6.6 86-7·4 86-8~5 86-87 86-8,6 86-8'9 86-90 86-91 86-9:2 86-9:3 86-9'9 86-11:)0 86-103 86-107 86-110 iotal Dry Ovary Weight (g) Sample a Sample b Sample c 1.6137 1.7968 1.6224 2.6649 1. 3212 1.0642 1.0485 1.1448 0.6912 0.6680 1.2961 5.0308 3.2729 3.5211 3.7196 3.2905 2.8967 2.2868 0.8488 4.2037 2.1033 2.8450 2.4860 1.1462 2.0159 2.1122 1.8504 3.0734 2.1863 1.6969 0.9892 1.0312 1.7264 4.7182 1.2032 4.0100 0.0307 0.0302 0.0303 0.0285 0.0303 0.0272 0.0302 0.0300 0.0311 0.0260 0.0277 0.0224 0.0253 0.0242 0.0268 0.0284 0.0285 0.0291 0.0267 0.0237 0.0249 0.0263 0.0279 0.0294 0.0294 0.0307 0.0286 0.0253 0.0289 0.0308 0.0294 0.0283 0.0298 0.0282 0.0304 0.0257 0.0304 0.0312 0.0316 0.0294 0.0296 0.0276 0.0295 0.0312 0.0307 0.0252 0.0272 0.0237 0.0270 0.0240 0.0284 0.0256 0.0280 0.0286 0.0266 0.0245 0.0257 0.0262 0.0292 0.0283 0.0301 0.0299 0.0302 0.0245 0.0291 0.0302 0.0275 0.0285 0.0303 0.0293 0.0299 0.0249 0.0305 0.0290 0.0307 0.0292 0.0296 0.0273 0.0301 0.0304 0.0300 0.0251 0.0264 0.0225 0.0262 0.0243 0.0283 0.0268 0.0292 0.0286 0.0256 0.0245 0.0259 0.0273 0.0290 0.0276 0.0301 0.0299 0.0301 0.0247 0.0296 0.0306 0.0290 0.0285 0.0299 0.0280 0.0298 0.0249 69 Append:lx 5. Volumetric fecundity data: estimates of total fecundity using individual subsamples, calculated from F = nV (see Methods and Materials): aeans of the individual estimates: and standard deviations (SD) of the individual estiaates. Fecundity Estimate Fish I.D. Number 85-1 85-2 85-4 85-5 85-6 85-7 85-8 85-9 85-13 85-15 85-16 85-17 85-18 85-19 85-20 85-22 85-24 85-25 85-26 85-27 85-29 85-32 85-33 85-35 85-37 85-39 85-40 86-3 86-4 86-5 86-6 86-9 86-12 86-14 86-15 86-18 86-19 86-20 86-21 86-22 86-23 Sample a 14 26 13 14 13 11 33 23 14 56 25 24 7 51 25 9 16 17 13 9 12 10 23 14 11 18 17 14 18 18 22 27 19 20 8 11 21 20 11 31 14 741 368 555 973 629 150 804 540 080 534 740 596 875 047 223 631 155 261 738 647 611 951 536 494 107 753 261 749 315 920 142 119 186 868 636 557 502 577 730 493 415 Sample b 15 27 15 15 15 11 36 23 14 63 26 26 8 52 24 9 15 17 14 9 12 11 23 15 11 19 17 14 18 18 22 27 19 20 8 11 20 20 12 31 14 020 263 356 812 222 331 180 276 840 406 520 098 435 203 640 901 840 633 029 293 002 154 587 206 542 646 112 620 482 705 533 775 765 979 693 739 534 235 121 820 136 Sample c 15 27 14 16 14 11 35 24 15 60 27 25 8 53 25 9 16 16 13 10 12 11 24 15 11 18 17 14 18 19 22 28 19 20 8 11 21 19 11 32 14 485 960 861 574 249 620 100 156 360 746 105 597 458 690 223 708 785 777 608 207 573 120 566 629 716 565 893 706 926 307 707 067 403 591 978 739 055 950 691 229 927 Mean 15 27 14 15 14 11 35 23 14 60 26 25 8 52 25 9 16 17 13 9 12 11 23 15 11 18 17 14 18 18 22 27 19 20 8 11 21 20 11 31 14 082 197 591 786 367 367 028 657 760 229 455 430 256 313 029 747 260 224 792 715 395 075 896 110 455 988 422 692 574 977 461 653 452 813 769 678 030 254 847 848 493 SO 376 798 930 800 803 237 1190 452 644 3465 685 765 330 1325 337 139 481 429 216 461 341 109 580 574 314 578 415 66 316 305 290 485 293 200 183 105 484 314 238 369 401 70 Appendix 5 continued. ----------------------------------------------------------Fecundity Estimate Fish 1.0. ------------------------------------Sa.ple a Sa.ple b Sa.ple c Mean Number SD ----------------------------------------------------------86-26 86-27 86-28 86-30 86-31 86-34 86-38 86-39 86-40 86-42 86-46 86-48 86-49 86-50 86-51 86-52 86-53 86-55 86-58 86-59 86-60 86-61 86-62 86-63 86-65 86-68 86-74 86-85 86-87 86-88 86-89 86-90 86-91 86-92 86-93 86-94 86-99 86-100 86-101 86-103 86-107 86-110 11 38 12 15 18 26 18 13 10 11 12 11 8 8 16 33 25 31 31 22 24 17 7 40 13 21 22 11 20 16 19 22 19 19 9 13 12 13 25 40 11 32 920 966 323 822 424 018 354 720 613 534 408 826 201 467 676 748 498 218 115 781 081 249 540 792 756 414 918 910 460 002 780 733 872 941 040 625 830 125 354 101 377 884 12 38 12 16 19 28 18 13 10 11 12 11 8 8 16 35 27 30 32 21 23 16 7 41 14 20 24 11 21 15 20 22 20 20 7 13 13 13 24 41 11 31 040 524 369 362 649 038 768 400 647 308 188 720 140 735 280 578 342 315 258 255 612 515 337 114 842 023 583 584 142 435 873 528 907 094 970 764 414 440 736 210 230 448 11 40 12 15 19 26 20 13 10 11 13 11 7 8 16 34 25 31 32 22 25 17 7 41 14 22 24 11 21 15 19 22 20 20 8 13 13 13 25 38 10 31 920 442 462 444 110 906 148 480 782 696 332 508 856 491 500 358 498 734 766 127 299 029 337 435 552 433 980 656 142 561 780 528 838 145 683 764 057 388 509 199 900 592 11 39 12 15 19 26 19 13 10 11 12 11 8 8 16 34 26 31 32 22 24 16 7 41 14 21 24 11 20 15 20 22 20 20 8 13 13 13 25 39 11 31 960 311 385 876 061 987 090 533 681 513 643 684 066 564 485 561 112 089 046 054 331 931 405 114 384 290 160 717 915 666 144 596 539 060 564 718 100 318 200 836 169 975 69 1005 71 461 614 1012 939 167 89 195 607 162 184 148 198 932 1065 718 846 766 871 377 117 321 562 1210 1094 171 394 298 631 118 579 106 544 81 294 169 409 1523 244 791 ----------------------------------------------------------- 71 Appendix 6. Grevimetric fecundity dete: estimetes of tote1 fecundity using individuel subsemples. celculated from F = nW/w (see Methods end Materials); means of the individual estimates; and standard deviations (SO) of the individual estimetea. Fecundity Estimate Fish I.D. Number 85-1 85-2 85-4 85-5 85-6 85-7 85-8 85-9 85-13 85-15 85-16 85-17 85-18 85-19 85-20 85-22 85-24 85-25 85-26 85-27 85-29 85-32 85-33 85-35 85-37 85-39 85-40 86-3 86-4 86-5 86-6 86-9 86-12 86-14 86-15 86-18 86-19 86-20 86-21 86-22 86-23 Semple e 16 31 15 17 15 12 37 24 16 63 28 28 8 54 26 10 17 18 15 10 13 11 25 16 12 20 19 15 19 20 24 30 20 22 9 12 23 20 13 33 16 792 290 526 924 118 229 786 329 550 110 339 458 885 557 652 930 922 256 098 736 803 710 973 054 430 365 485 797 728 454 500 630 683 750 744 702 133 444 070 964 128 Semple b 16 31 16 18 16 12 38 24 17 67 28 28 9 54 27 10 17 18 15 10 14 11 25 16 12 20 19 15 19 20 25 30 20 22 '3 12 22 20 13 34 15 441 128 358 675 141 919 517 759 268 007 992 810 756 692 007 634 516 825 418 823 105 967 945 348 634 982 513 857 908 093 107 948 793 871 809 404 092 578 339 429 354 Semple c 16 31 16 19 15 13 38 25 17 65 29 28 9 54 27 10 17 18 15 11 15 11 26 16 12 21 19 15 19 20 25 30 20 23 9 12 22 20 13 33 15 679 823 000 254 862 119 092 055 071 506 528 786 610 458 276 662 759 551 368 044 671 891 588 802 550 233 480 461 685 544 567 654 554 297 900 404 403 495 150 768 752 Meen 16 31 15 18 15 12 38 24 16 65 28 28 9 54 26 10 17 18 15 10 14 11 26 16 12 20 19 15 19 20 25 30 20 22 9 12 22 20 13 34 15 638 414 961 618 707 756 132 714 963 207 953 685 417 569 978 742 732 544 294 868 526 856 169 402 538 860 493 705 774 364 058 744 677 973 818 503 543 506 186 054 745 SD 179 364 417 667 529 467 367 365 371 1966 595 197 466 117 313 163 204 284 172 159 1003 132 364 377 103 447 18 213 118 238 535 177 120 287 78 172 534 68 138 340 387 ----------------------------------------------------------- 72 Appendix 6 continued. Fecundity Estimate Fish r .. D. Number 86-26 86-27 86-28 86-30 86-31 86-34 86-39 86-40 86-42 86-48 86-49 86-50 86-51 86-52 86-53 86-55 86-58 86-59 86-60 86-61 86-62 86-63 86-65 86-68 86-74 86-85 86-87 86-88 86-89 86-90 86-91 86-92 86-93 86-99 86-100 86-103 86-107 86-110 Sample a 13 40 13 17 20 30 14 12 12 12 8 8 17 37 30 35 34 24 26 18 8 45 16 24 25 12 22 17 22 26 21 21 9 14 14 42 12 35 297 799 929 433 133 109 956 285 360 784 979 915 734 282 401 211 004 215 121 467 266 052 049 988 751 827 628 476 257 968 787 542 387 430 483 330 270 731 Sample b 13 40 14 17 20 31 14 12 12 12 9 9 17 37 30 34 33 25 26 17 8 43 16 23 26 12 22 17 22 27 22 22 8 14 14 41 12 35 525 733 120 450 588 453 953 146 404 182 028 490 631 147 547 478 267 064 070 991 073 924 777 455 392 961 838 307 242 598 765 138 849 980 586 868 314 269 Sample c 13 40 14 17 20 30 15 12 12 12 8 9 18 37 29 35 33 24 26 18 8 44 16 25 27 13 22 17 21 27 22 21 9 14 14 40 11 35 532 879 179 720 610 391 042 435 575 277 917 262 411 787 356 646 910 924 785 550 389 268 323 219 003 372 838 449 147 374 306 904 142 582 724 610 992 430 Mean 13 40 14 17 20 30 14 12 12 12 8 9 17 37 30 35 33 24 26 18 8 44 16 24 26 13 22 17 21 27 22 21 9 14 14 41 12 35 452 804 076 534 444 651 984 289 446 414 975 222 925 405 101 111 727 735 325 336 242 415 383 554 382 053 768 411 882 314 286 862 126 664 598 603 192 477 SD 134 73 131 161 270 709 51 145 114 324 56 289 424 337 649 590 401 455 398 302 159 578 368 959 626 284 122 90 636 319 489 301 270 284 121 890 175 235