36 CALIFORNIA COOPERATIVE OCEANIC FISHERIES INVESTIGATIONSfurther offshore (Blackburn 1941, 1950 a). There isno reliable information about the vertical distributionof E. australis in the sea. It is found frequently inthe stomachs of large scombriform fish hooked at thesea surface (Blackburn 1950 a), but not at all in thestomachs of Neoplatycephalus macrodon, which is oneof the most abundant and active demersal fishes ofcontinental shelf waters over 50 m (Fairbridge1951) ; this suggests that it is much commoner nearthe surface than near the bottom, in those waters. Inshallower waters, such as 20 m, echograms show thatthe species can occur from top to bottom (Figure 2).Charts of surface temperature and salinity fromvarious sources (Xchott 1935; U.X. Navy HydrographicOffice 1944; U.S.S.R. Ministry of Defense1953 ; Muromtsev 1958 ; Rochford 1962) have beencompared with the general spatial distribution ofE. australis (Figure 1) and the seasonal changes thatoccur in it. This showed that the species toleratestemperatures at least from 10 to 21" C.; it may encountertemperatures up to 25" C. if it occurs insurface waters as far north in summer as it does inwinter, but this is not certain ; it probably encounters9" C. in certain inshore waters in winter, but thisalso is not certain. The same comparison shows atolerance of salinities from about 35.1 to 35.8%0, butdata obtained from various inshore waters, at timesand places where anchovies occurred, indicate thatthe range is actually from about 2 to 37%0 (Blackburn1950 a, Gippsland Lakes rivers; Rochford 1957, St.Vincent's Gulf). The range of breeding temperaturewas investigated by comparing seasonal temperaturedata (including those for bays and estuaries in variousvolumes of the Australian " Oceanographical StationList ") with information on spaxning places andseasons; this range is at least 14 to 20" C. and possiblyincludes 13" C.The above-mentioned temperature ranges for E.australis are almost the same as those for the Australiansardine, Sardinops neopilchardus, which hasan almost identical distribution along the Australiancoast (Blackburn 1960 and references). The sardineis not known to occur in waters of salinity belowabout 33.5%0, however, and thus it is not estuarine,whereas the anchovy is. Another difference in spatialdistribution between the two species is obvious whentheir relative abundances are compared within differentparts of their common total range of distribution(including saline bays and gulfs where bothspecies occur) j to the north of about 37" S. latitudethe sardine is more abundant than the anchovy (shownmainly by differences in numbers of larvae, as thereare no good data for eggs or adults of the anchovy;Blackburn 1941) ; to the south of 37" S. the anchovyis more abundant than the sardine (shown by differencesin numbers of eggs and larvae, and generalscarcity of adult sardines ; Blackburn 1950 a, 1950 b,1957).INTRASPECIFIC POPULATIONSBlackburn (1950 a) presented and analyzed thetotal vertebral counts, including urostyle, of 5,804 E.australis from 21 localities around the Australiancoast. Analysis of variance indicated that significantdifferences existed between vertebral count means forcertain localities, and the variability was furtherinvestigated by the graphical method of Hubbs andPerlmutter (1942).Figures 3 and 4, from Blackburn (1950 a), showthe result. The localities listed in Figure 3 appearin the order in which they occur around the Australiancoast, from north-east (top) to south-west(bottom). With one exception (Gippsland Lakesrivers, discussed below) the means fall clearly intoeastern, southern, and western groups which weregiven subspecific names (E. a. australis, E. a. antipodum,and E. a. fraseri, respectively). The localitymeansin the eastern group show a cline with meansincreasing from north to south (28 to 36" S. latitude,43.1 to 44.0 vertebrae). Means in the western groupare numerically similar to those of the eastern groupat similar latitudes, and suggest a similar cline. Theexistence of clines, whether north-south or east-west,is doubtful in the means of the southern group ; thesemeans are all at least one vertebra higher (45.1 to45.8 vertebrae) than any of the means of the othertwo groups, and this difference is statistically highlysignificant. Some of the differences between meanswithin each group are significant.The Gippsland Lakes are situated on the east coastof the state of Victoria. As shown in Figure 5(left inset), this is a system of three rivers enteringa coastal lagoon or lake which is permanently open tothe sea. E. australis occurs in the sea outside the entrance,at the entrance, in the lake, and in the rivers(Blackburn 1950 a). Samples suEcient for vertebralcount study were available only from the entrance(salinity about 35.5%0) and the rivers (salinity about2%0), and these two-localities are distinguished inFigure 3. The river fish have a mean about two-thirdsof a vertebra lower than the mean for the entrance, ahighly significant difference. This finding resemblesthat of Hubbs (1925) for Engraulis mordax in SanFrancisco Bay, <strong>California</strong>.I.-CQOWAITA R BO.__Qczzz: E; ",. -LUACWUU1. nJ*uuo* Iuw. NSW DO. (00J m L NLW UWAOOUOA1. NSW (01-a NLW suMUC NSW 7X0IrunLomr)rc. ne anOI~LIYO L IYP. VIC mrrmr*rm vh a00murmiwa IL smau.ar .,T U-87 UCLm TAL XI0-aru m~ ~ ~ ~ C L . R W X ~ I I I I I TAL ~.oMco*r TAL (01COmLU Io.m I w L R V I a7swmn WA n-MI1 0 . I U U Y U Y . 7 BYtlRU.FIGURE 3. Summary of data on vertebral counts of E. australis. Numbenat the left are numbers of counts from different localities. Thehorizontal line for each sample is the range, the vertical cross-bar isthe mean, the thickened portion of the line is one standard deviationon each side of the mean, and the hollow rectangle is twice thestandard error on each side of the mean (from Blackburn 19500).
REPORTS VOLUME XI, 1 JULY 1963 TO 30 JUNE 1966 37FIGURE 4. Approximate disposition of subspecies of E. ausfralis in Australian waters, based on data in Figure 3. Arrows indicate clines in meanvertebra numbers (from Blackburn 1950a).Although these various differences in mean vertebranumber may be phenotypic rather than genotypic,their size and geographical distribution (and, for somewell-studied localities, their consistency over manyyears) indicate that E. australis is a complex of differentlocal, at least partly segregated, groups. Thereare three major groups (two of them morphologicallysimilar, but separated geographically by a third whichis morphologically different) ; within these, minor,less well differentiated, groups occur.GROWTHGrowth in length was investigated by scale-readingand inspection of length frequency distributions(Blackburn 1950 a). These studies were reasonablyadequate only for Victorian fish (members of thesouthern subspecies), for which about 21,000 lengthmeasurements, and scales from 450 fish, were availableand useful. Samples from the other five states yieldedscales from 314 fish, and about 12,000 lengths whichwere obtained too irregularly to be useful for lengthfrequencystudies. The deciduous nature of the scaleslimited the amount of scale material available, especiallyfor small fish.About three scales, all taken from the flank in theregion of the dorsal fin, were usually read for eachfish. The appearance of scales from this region isshown in Figures 6, 7, 8, and 9 (from Blackburn
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FISH E RYThe California anchovy fis
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CONTENTSI. Review of Activities Pag