390 
Abstract— Gonadal morphology and 
reproductive biology of the Black 
Anglerfish ( Lophius budegassa ) were 
studied by examining 4410 speci- 
mens collected between June 2007 
and December 2010 in the north- 
western Mediterranean Sea. Ovaries 
and testes presented traits common 
among fishes of the order Lophi- 
iformes. Spawning occurred between 
November and March. Size at first 
maturity (L 50 ) was 33.4 cm in total 
length (TL) for males and 48.2 cm 
TL for females. Black Anglerfish is 
a total spawner with group-synchro- 
nous oocyte development and de- 
terminate fecundity. Fecundity val- 
ues ranged from 87,569 to 398,986 
oocytes, and mean potential fecun- 
dity was estimated at 78,929 (stan- 
dard error of the mean [SE] 13,648) 
oocytes per kilogram of mature fe- 
male. This study provides the first 
description of the presence of 2-3 
eggs sharing the same chamber and 
a semicystic type of spermatogen- 
esis for Black Anglerfish. This new 
information allows for a better un- 
derstanding of Black Anglerfish re- 
production — knowledge that will be 
useful for the assessment and man- 
agement of this species. 
Manuscript submitted 19 October 2012. 
Manuscript accepted 19 September 2013. 
Fish. Bull. 111:390-401 (2013). 
doi: 10.7755/FB. 11 1.4.8 
The views and opinions expressed or 
implied in this article are those of the 
author (or authors) and do not necesarily 
reflect the position of the National 
Marine Fisheries Service, NOAA. 
Reproductive biology of Black Anglerfish 
C Lophius budegassa) in the northwestern 
Mediterranean Sea 
Ana I. Colmenero (contact author) 
Victor M. Tuset 
Laura Recasens 
Pilar Sanchez 
Email address for contact author: colmenero@icm.csic.es 
Institut de Ciencies del Mar (ICM) 
Consejo Superior de Investigaciones Cientificas (CSIC) 
Passeig Maritim de la Barceloneta 37-49 
08003 Barcelona, Spain 
Lophius, a genus commonly known as 
anglerfishes or monkfishes, includes 
7 species broadly distributed and ex- 
ploited worldwide. Most of these spe- 
cies inhabit the northwestern Atlan- 
tic, as do Goosefish ( Lophius ameri- 
canus ) and Blackfin Goosefish (L. 
gastropliysus ), or the northeastern 
Atlantic, as do Cape Monk (L. vome- 
rinus), Shortspine African Angler (L. 
vaillanti), Black Anglerfish (L. bude- 
gassa), and White Anglerfish (L. pis- 
catorius), although Black Anglerfish 
and White Anglerfish also live in 
the Mediterranean Sea and Yellow 
Goosefish (L. litulon ) can be found 
only in the northwestern Pacific 
(Farina et al., 2008). In the past, spe- 
cies of Lophius have been captured 
as bycatch in mixed fisheries, but 
an increase in their economic value, 
together with the overexploitation 
of other groundfish species, has led 
to the development of targeted an- 
glerfish fisheries (Hislop et al., 2001). 
In the northwestern Mediterranean 
Sea, landings of Black Anglerfish 
and White Anglerfish have accumu- 
lated to just over 6000 metric tons 
during the last 10 years. 1 The scarce 
reproductive information available 
for these species does not allow for a 
1 Tudo Vila, P. 2012. Unpubl. data. 
Directorate of Fishing and Maritime Af- 
faires, Government of Catalonia, Avin- 
guda Diagonal, 523-525, 08029 Barce- 
lona, Spain. 
proper assessment or informed man- 
agement of anglerfish fisheries. 
This study focuses on Black An- 
glerfish, a demersal fish distributed 
along the Mediterranean Sea, as well 
as the northeastern Atlantic from 
the British Isles to Senegal (Caruso, 
1986). This species is found over the 
continental shelf and upper slope at 
depths of up to 800 m and inhabits 
sandy, muddy, and rocky bottoms 
(Carlucci et al., 2009). They occupy 
the water column as eggs and larvae, 
and then they shift to a benthic exis- 
tence as juveniles and adults (Farina 
et al., 2008). This species co-occurs 
with White Anglerfish over all its 
bathymetric range, although White 
Anglerfish has a deeper distribu- 
tion that reaches to depths >1000 m 
(Afonso-Dias and Hislop, 1996). De- 
spite the overlapping distributions of 
these species, Colmenero et al. (2010) 
concluded that no ecological compe- 
tition exists between these species 
because of a temporal segregation 
in their biorhythms; Black Angler- 
fish is more active at nighttime, and 
White Anglerfish is more active dur- 
ing daytime. 
Most studies of these species 
have been undertaken in northeast- 
ern Atlantic waters, and they often 
have dealt with age and growth 
(Dupouy et al., 1984; Landa et al., 
2008a; Woodroffe et al., 2003), feed- 
ing habits (Crozier, 1985; Laurenson 
