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Fishery Bulletin 1 1 1 (4) 
Table 1 
Macroscopic and microscopic description of the 5 maturity phases in the reproductive cycle of male and female Black An- 
glerfish (Lophius budegassa ) collected from the northwestern Mediterranean Sea between June 2007 and December 2010; 
adapted from Afonso-Dias and Hislop (1996) and Brown-Peterson et al. (2011). 
Phase 
Males 
Females 
Immature (I) 
Testes are long, narrow, and tubular 
shaped. They are translucent with no 
visible vascularization. The medial semi- 
niferous duct is distinct. Only spermato- 
gonia and primary spermatocytes are 
present. 
Ovaries are very narrow, thin, and flattened-tube 
shaped. They are translucent; no oocyte clusters 
visible and minimal vascularization. Only oogonia 
and primary growth oocytes are present. 
Developing/ 
Regenerating (II) 
Testes are small with visible blood ves- 
sels around the seminal duct. Spermato- 
gonias, primary and secondary spermato- 
cytes are predominant. Spermatids are 
scarce. 
Ovaries are small. Still no noticeable individual oo- 
cyte clusters. They acquire a cream color and vas- 
cularization is visible. Only oogonia and primary 
growth oocytes are present. 
Spawning capable (III) 
Testes increase in length and width. They 
have a firm texture and cream color. Sem- 
inal duct is highly vascularized. Germ 
cells at all stages of spermatogenesis are 
present. Spermatids are predominant 
with a lot of spermatozoa in the lumen of 
the sperm duct. 
Ovaries increase in width and length. They have a 
light orange color, and blood vessels are prominent. 
The edges of the ovaries start to curl and they oc- 
cupy a larger proportion of the body cavity. A mucus 
matrix starts to develop. Primary growth, cortical 
alveolar, and primary and secondary vitellogenic oo- 
cytes are present. 
Actively spawning (IV) 
Testes are large and firm and have a 
creamy coloration. Large amounts of 
sperm produced when testes are dissect- 
ed. Abundant quantities of spermatozoa 
are present in the seminiferous tubules. 
Ovaries are extremely long and wide and occupy 
most of the body cavity. The color of the oocytes is or- 
ange, and they are visible macroscopically. Ovaries 
are characterized by the presence of large hyaline 
oocyte clusters enclosed in a transparent gelatinous 
matrix that is completely developed. High vascular- 
ization is present. Oocytes are in tertiary vitellogen- 
esis, migratory nucleus and hydration. 
Regressing (V) 
Testes are small, flaccid, and have brown 
or red areas in their beige surface. They 
are still highly vascularized. Sperm and 
residual spermatozoa can be found in the 
lumina of the sperm duct. Spermatogonia 
are present in the testes cortex. 
Ovaries are flaccid and highly vascularized and of- 
ten have longitudinal striations. Their color is dark 
pink or red. Atresia and postovulatory follicles, 
together with primary growth stages, are present. 
Cortical alveolar, primary and secondary vitellogen- 
esis can be found. 
and from 0°32.922'E to 3°35.718'E between June 2007 
and December 2010. Specimens were collected onboard 
commercial trawl fishery vessels at depths of 20-600 
m and identified according to Caruso (1986). The trawl 
fleet belonged to the ports of Roses, Blanes, Arenys de 
Mar, Vilanova i la Geltru, and Sant Carles de la Rapita 
(Fig. 1). For this study, 4410 specimens were measured 
to the nearest centimeter in total length (TL), weighed 
to the nearest gram in total weight (TW) and gutted 
weight (GW), and measured with an accuracy of 0.01 g 
in gonad weight (GNW) and liver weight (LW). 
Macroscopic and histological description of gonads 
Of the total number of specimens, 3562 fish had gonads 
removed and their sex was determined, and they were 
assigned macroscopically to a gonadal stage on the basis 
of a scale with 5 maturity phases that were described 
in previous studies: immature (phase I), developing or 
regenerating (phase II), spawning capable (phase III), 
actively spawning (phase IV), and regressing (phase V) 
(Afonso-Dias and Hislop, 1996; Brown-Peterson et ah, 
2011) (Table 1). Sex was easily assessed macroscopical- 
ly in mature individuals. However, gonads from small 
individuals (approximately <20 cm TL) were indistin- 
guishable macroscopically because ovaries and testes 
were small, translucent, and string-like. Fish that were 
too small to determine their sex or assign to a gonadal 
phase were classified as indeterminate. 
To corroborate the macroscopic classification of 
some unclear and undetermined gonads, 372 speci- 
mens were histologically examined. They were fixed in 
10% buffered formalin solution before they were dehy- 
drated and embedded in a methacrylate polymer resin. 
