Fitzhugh et al.: Size- and age-dependence in batch spawning 
419 
Table 4 
Patterns of spawning duration by age or size based on a review of spawning season duration within the scientific literature on 
fish reproduction. 
Spawning 
Duration 
Species 
Reference(s) 
Increases with 
Atkerina boyeri (sand smelt, Atherinidae) 
Tomasini et al., 1996; Tomasini and 
size or age 
Laugier, 2002 
Clupea bentincki (Chilean herring, Clupeidae) 
Cubillos and Claramunt, 2009 
Cynoscion nebulosus (spotted seatrout, Sciaenidae) 
Lowerre-Barbieri et al., 2009 
Engraulis encrasicolus (European anchovy, Engraulidae) 
Millan, 1999 
Engraulis mordax (northern anchovy, Engraulidae) 
Parrish et al., 1986 
Engraulis ringens (Peruvian anchoveta, Engraulidae) 
Claramunt et al., 2007; Cubillos and 
Claramunt, 2009 
Gadus morhua (Atlantic cod, Gadidae) 
Kjesbu et al., 1996; Lawson and Rose, 
2000 
Genyonemus lineatus (white croaker, Sciaenidae) 
Love et al., 1984 
Hemiramphus brasiliensis (ballyhoo, Hemiramphidae) 
McBride and Thurman, 2003 
Lepomis gibbosus (pumpkinseed, Centrarchidae) 
Fox and Crivelli, 1998 
Leuresthes tenuis (California grunion, Atherinidae) 
Clark, 1925 
Lutjanus carponotatus (stripey sea perch, Lutjanidae) 
Kritzer, 2004 
Melanogrammus aeglefinus (haddock, Gadidae) 
Trippel and Neil, 2004 
Merluccius hubbsi (Argentine hake, Merlucciidae) 
Macchi et al., 2004 
Sardina caerulea (Califorinia sardine, Clupeidae) 
Clark, 1934 
Sardina pilchardus (true sardine, Clupeidae) 
Zwolinski et al., 2001 
Sardina pilchardus sardina (Mediterranean sardine, Clupeidae) 
Ganias et al., 2003b 
Sardinops ocellatus (southern African pilchard, Clupeidae) 
LeClus, 1989 
Sardinops sagax (Pacific sardine, Clupeidae) 
Claramunt and Herrera, 1994 
Scomber japonicus (Pacific chub mackerel, Scombridae) 
Dickerson et al., 1992; Knaggs and 
Parrish 1 ; MacCall et al., 1980 
Seriphus politus (queenfish, Sciaenidae) 
DeMartini and Fountain, 1981 
Solea solea (sole, Soleidae) 
Witthames 2 
Trachurus symmetricus (jack mackerel, Carangidae) 
Wine and Knaggs 3 
No change noted 
Clupea harengus (Atlantic herring, Clupeidae 
Lambert, 1987 
overall, but larger 
Clupea pallasi (Pacific herring, Clupeidae) 
Ware and Tanasichuk, 1989 
fish develop eggs 
Cynoscion regalis (weakfish, Sciaenidae) 
Lowerre-Barbieri et al., 1996 
and/or spawn 
Scomber scombrus (Atlantic mackerel, Scombridae) 
Priede and Watson, 1993 
earlier 
Trachurus trachurus (horse mackerel, Carangidae) 
Abaunza et al., 2003 
1 Knaggs, E. H., and R. H. Parrish. 1973. Maturation and growth of Pacific mackerel Scomber japonicus Houttuyn. Marine Resources Tech. 
Rept. 3, 19 p. Calif. Dept. Fish and Game, Long Beach, CA. 
2 Witthames, P. R. 
2003. Methods to assess maturity and realized fecundity illustrated by studies on Dover sole Solea solea. In Report of the 
working group of modern approaches to assess maturity and fecundity of warm- and cold-water fish and squids; 4-7 September 2001, Bergen, 
Norway (O. S. Kjesbu, J. R. Hunter and P. R. Witthames, eds.), Fisken og havet. no. 12-2003, 
p. 125-138. Inst. Mar Res., Bergen, Norway. 
3 Wine, V. L., and E. 
H. Knaggs. 1975. Maturation and growth of jack mackerel, Trachurus symmetricus. Mar. Resources Tech. Rept 32, p. 
1-25. Calif Dept. Fish and Game, Long Beach, CA, 
also found that older fish had greater annual fecundity, 
and less frequent spawning over a longer season. In 
other cases, mediating factors, such as temperature or 
productivity, may have explained an anomalous size- 
based trend. Two articles on anchovies and a sardine 
reported lower spawning frequency among larger-size 
fish of stocks in shelf waters that were geographically 
separated from smaller fish in warmer coastal waters 
(Cubillos et ah, 2007; Dimlich et ah, 2009). Elsewhere, 
one of the anchovy species, Peruvian anchoveta ( Engrau - 
lis ringens), was noted to have a positive relationship of 
size with both spawning frequency (Claramunt et al., 
2004, 2007) and duration (Claramunt et al., 2007; Cubil- 
los and Claramunt, 2009). From life-history theory, we 
would expect older fishes to invest relatively more energy 
into reproduction (Stearns, 1992); however, increased 
investment could manifest in various ways (e.g., egg 
quality rather than quantity; Berkeley et al., 2004a, 
2004b). Empirical observations, as reviewed here, were 
necessary to elucidate patterns in the annual number of 
batches. The common assumption of age or size invari- 
ance would appear to be generally wrong. 
