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Fishery Bulletin 95(4), 1997 
Table 1 
Histological criteria developed by MARMAP (Charleston, SC) to determine reproductive stage in red porgy, Pagrus pagrus (see 
D’Ancona, 1949, 1950; Wallace and Selman, 1981; Alekseev, 1982, 1983; Hunter etal., 1986; Sadovy and Shapiro, 1987; Matsuyama 
et al., 1988; West, 1990; Roumillat and Waltz 7 ). 
Reproductive state 
Male 
Female 
Immature (virgin) 
No primary males found. Juveniles were either fe- 
males or, infrequently, simultaneous or transitional 
(see below). 
Previtellogenic oocytes only; no evidence of 
atresia. In comparison with resting female, 
most previtellogenic oocytes <80 pm, area of 
transverse section of ovary is smaller, lamel- 
lae lack muscle and connective tissue bundles 
and are not as elongate, germinal epithelium 
along margin of lamellae is thicker, ovarian 
wall is thinner. 
Developing 
Development of cysts containing primary and sec- 
ondary spermatocytes through some accumulation 
of spermatozoa in lobular lumina and dorsomedial 
sinuses. 
Oocytes undergoing cortical granule (alveoli) 
formation through nucleus migration and 
partial coalescence of yolk globules. 
Running and ripe 
Predominance of spermatozoa in lobules and 
dorsomedial sinuses; little or no occurrence of sper- 
matogenesis. 
Completion of yolk coalescence and hydration 
in most advanced oocytes. Zona radiata becomes 
thin. Postovulatory follicles sometimes present. 
Developing, recent spawn 
Not assessed. 
Developing stage as described above as well 
as presence of postovulatory follicles. 
Spent 
No spermatogenesis; some residual spermatozoa 
in lobules and sinuses. 
More than 50% of vitellogenic oocytes with 
alpha- or beta-stage atresia. 
Resting 
Little or no spermatocyte development; empty lob- 
ules and sinuses. 
Previtellogenic oocytes only; traces of artresia. 
In comparison with immature female, most 
previtellogenic oocytes >80 pm, area of trans- 
verse section of ovary is larger, lamellae have 
muscle and connective tissue bundles, lamel- 
lae are more elongate and convoluted, germi- 
nal epithelium along margin of lamellae is 
thinner, ovarian wall is thicker. 
Mature specimen, 
stage unknown 
Mature, but inadequate quantity of tissue or post- 
mortem histolysis prevent further assessment of 
reproductive stage. 
Mature, but inadequate quantity of tissue or 
postmortem histolysis prevent further assess- 
ment of reproductive stage. 
Simultaneous (bisexual) 
Presence of distinct ovarian and testicular regions in approximately equal amounts and of the same 
reproductive state. This gonad structure was infrequently observed in both juvenile and adult fish. 
Transitional 
Ventrolateral proliferation of active testicular tissue (spermatogonia through spermatozoa) along 
the outer surface of the ovarian wall in spent or resting ovary (functional protogyny) or immature 
ovary (juvenile protogyny). As testicular tissue envelopes regressing ovary, ovary collapses laterally 
and sperm sinuses form within former ovarian wall. 
1 Roumillat, W. A., and C. W. Waltz. 1993. Biology of the red porgy, Pagrus pagrus, from the southeastern United States. MARMAP Final Data 
Report, South Carolina Department of Natural Resources, Charleston, SC, 38 p. 
rate (i.e. the negative relation between L ^ and k). 
However, k was highest for the 1979-81 period, when 
was also still relatively high. 
Reproduction 
Our examination of 4,293 gonads (n=l,397, 1979- 
81;n=727, 1988-90; n=2, 169, 1991-94) revealed that 
sexual transition was occurring at smaller sizes in 
the later periods. There was a significant increase 
(P<0.001) in the number of males with time (Table 
4). However, in 1988-90 and in 1991-94, the propor- 
tion of males to the total number of fish sexed was 
significantly greater at smaller sizes than during 
1979-81 (Table 4). At 301-350 mm TL, male red 
porgy made up 24% of the fish that were sexed dur- 
