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Received: 12 July 2018 Accepted: 18 February 2019

DOI: 10.1111/jfb.13937

FISH
SYMPOSIUM SPECIAL ISSUE REGULAR PAPER

Can we consider the stocks of alfonsinos Beryx splendens and


Beryx decadactylus from the Azores a discrete fishery
management unit?
Régis V. S. Santos1,2,3 | Ana M. Novoa-Pabon2,4 | Hélder M. Silva1,2,3,4 |
Mário R. Pinho1,2,3,4

1
IMAR Institute of Marine Research,
University of the Azores, Horta, Portugal This paper summarises the detailed information on catch, fork length, weight, sex and maturity
2
Okeanos R&D Centre, University of the of alfonsinos Beryx splendens and Beryx decadactylus collected from targeted surveys in Azores
Azores, Horta, Portugal waters, complemented by fishery data, over the past c. 20 years. To date, it was not possible to
3
MARE Marine and Environmental Sciences define if the Azorean component of the population can be considered as a discrete local man-
Centre, University of the Azores, Horta,
agement unit. The reason for this is the lack of available information for these resources cover-
Portugal
4
ing the entire spatial distribution of the species in the North Atlantic Ocean. Additionally, there
Faculty of Science and Technology,
Department Oceanography and Fisheries, are some conflicts between the different genetic results available, poor understanding about the
University of the Azores, Horta, Portugal species movements and data from the Azores show some discrepancies in aspects of reproduc-
Correspondence tion. Consequently, there is no analytical assessment for this resource and the stock of Beryx
Régis V. S. Santos, IMAR Institute of Marine spp. is currently managed based on the precautionary approach. Outputs of recent analyses are
Research, University of the Azores, 9901-862
presented in this study and possible assumptions and strategies for the assessment are
Horta, Portugal.
Email: regisvinicius@gmail.com discussed.
Funding information
This study was funded by the Azorean KEYWORDS
Government under the Data Collection
assessment, Beryx decadactylus, Beryx splendens, commercial fish, FMU, management
Framework (Council Regulation (EC) No
199/2008) of European Commission and the
DEMERSAIS project. Régis Santos was funded
by the IMAR Instituto do Mar, through a Post-
doc fellowship (ref. IMAR/DEMERSAIS/001-
2018). Ana Novoa-Pabon was funded by a FCT
Ph.D. fellowship (ref. SFRH/BD/124720/2016).

1 | I N T RO D UC T I O N management measures adopted on these units (Uriarte et al., 2014).


This involves good definition of the stocks being exploited, their spa-
A fishery management unit (FMU) is a discrete area of the ecosystem tial distribution and biology.
and its component fisheries that are the focus for management under Alfonsinos fisheries are found in almost all seas where moder-
an ecosystem approach to fisheries management (Staples et al., 2014). ately deep waters exist, in both tropical and temperate latitudes,
Thus, an FMU can be a specific type of fishing (e.g., longline fishery) or except in the North-east Pacific Ocean (FAO, 2016). In the North
a resource (e.g., alfonsino fishery) or a geographic area. In order to Atlantic Ocean, two Beryx Cuvier 1829 species, splendid alfonsino
implement a sustainable fisheries management strategy, management Beryx splendens Lowe 1834 and alfonsino Beryx decadactylus Cuvier
units should ensure the matching of biologically relevant processes 1829, commonly referred to as alfonsinos, are currently caught by
and management actions (Reiss et al., 2009). In other words, the man- demersal trawl and longline mixed fisheries targeting deep-water
agement units must comprise, within their limitations, much of the life species. Both species are benthopelagic as adults but have epipe-
cycle of the species in the community being managed, in order to lagic eggs and larvae and pelagic juveniles (Isidro, 1996;
ensure that the recovery of the resources is associated to the Mundy, 1990).

J Fish Biol. 2019;1–8. wileyonlinelibrary.com/journal/jfb © 2019 The Fisheries Society of the British Isles 1
2
FISH SANTOS ET AL.

Beryx splendens normally lives close to, or within 5–20 m, of the Ocean (ICES area 10). The archipelago consists of nine volcanic
rocky bottoms of the upper slope between depths of 200 and 1240 m islands forming three groups, running from west-northwest
but typically around seamounts and deep-water reefs in waters (WNW) to east-southeast (ESE) between 36  and 40 N latitude,
200–800 m deep (Maul, 1981). It is widely distributed and can be 24 and 32  W longitude (Figure 1). Most of the surface of the oce-
found in the deep-waters of the Atlantic, Indian and Pacific Oceans anic region around the Azores is abyssal plain with average
(FAO, 2016). The species inhabits the western Atlantic Ocean from depth > c. 4000 m. The major topographic feature is the Mid-
Nova Scotia to Brazil, the Corner Rise seamounts (35 230 2800 N 51 Atlantic Ridge that follows a sinuous course southward from Ice-
400 3900 W) and around other islands in the South Atlantic Ocean land to the Azores. Islands, seamounts and hydrothermal vent
(Alekseev et al., 1986). In the eastern Atlantic Ocean, B. splendens is fields are the prominent features along this deep-sea environment.
found from Iceland and Norway to South Africa and around the The land area of the archipelago is 2344 km2 and the marine exclu-
Azores, Madeira, Canaries, Saint Helena and Tristan da Cunha (FAO, sive economic zone (EEZ) is 948,439 km2 (Instituto Hidrográfico,
2016). Beryx decadactylus is also found along the Mid-Atlantic Ridge, 1981). The shallow areas (< 700 m), where the most fishing effort
north and south of the Azores, on rocky bottoms and normally lives
occurs, are very limited and with considerable discontinuity (Pinho,
between 200 and 900 m depth, most commonly found between
2003; Menezes et al., 2006). The marine topography is highly vari-
400 and 600 m (Maul, 1981, 1986).
able, characterised mainly by rocky bottoms where there is no con-
For most of the fisheries, the catches are reported under a single
tinental shelf and sedimentary areas are scarce (Martins, 1986).
category, as Beryx spp. and the proportions of each species in the
catches are not well known, as well as the distribution, reproductive
variables, age, growth and mortality rates. Because very little is known 2.2 | Sample collection
about stock structure of these species, the International Council for
Surveys were conducted annually each spring (usually from March–
the Exploration of the Sea (ICES) working group on the biology and
April to June) from 1995 to 2017 during cruises onboard the
assessment of deep-sea fisheries resources (WGDEEP) currently
R.V. Arquipélago. The survey follows a stratified random design and
assesses a single stock comprising both species in the North Atlantic
covers the Azores Archipelago around the islands, banks and major
Ocean (ICES, 2017). Detailed landings data by species are available
seamounts as far as 70 nm from the islands. Each sampling area is fur-
only for the Azores area (ICES Subarea 10a2; www.ices.dk/marine-
ther divided into depth strata with 50 m intervals (i.e., the first stratum
data/maps/Pages/default.aspx), where the landings of B. decadactylus
averaged 20% of the catches of both species (ICES, 2017). represents 0–50 m, the second 51–100 m and so on) down to 800 m

Over the past c. 20 years, surveys in the Azores archipelago, com- depth and for some pre-defined sets, down to 1200 m for abundance

plemented by fishery data from the European Commission's data col- estimation and ecological purposes. Each fishing set is laid in a way to

lection framework (DCF; EU, 2008), have recorded information such cross the different depth strata. So, each set is a vertical transect from
as catch, fork length (LF), mass, sex and maturity stage. However, it the surface to the bottom, covering whenever possible 24 depth
was not possible until now to define if the Azorean component of the strata (stations). On average, c. 510 stations (area–depth strata) have
Beryx spp. population could be considered as a management unit. The been sampled annually. The survey gear (bottom longline design for
reason for this is the lack of available information for these resources benthopelagic species) is very similar to the one used by the commer-
covering the entire spatial distribution of the species in the North cial fishery. The bait is chopped salted European pilchard Sardina pil-
Atlantic Ocean. Associated to this, there are some conflicts between chardus (Walbaum 1792). All fishes are tallied by species and strata,
the different genetic results available, poor understanding of the spe- measured and weighed. Detailed biological sampling by strata is made
cies’ movements and data from the Azores show some discrepancies based on a subsample of the total catch and information on LF, mass,
in relation to reproduction. Consequently, there is no analytical sex, otoliths, maturity stage, gonad and liver weight, are recorded.
assessment for this resource and hence the stock of Beryx spp. is cur- Some fishes also are sampled for genetics and heavy metals contami-
rently managed according to the precautionary approach (FAO, 1996). nation. Detailed information on survey design are given by (2003),
This study results from a recommendation of the ICES WGDEEP Menezes et al. (2006) and ICES (2010).
in 2018, with the objective of examining the population structure of Survey data were complemented by fisheries data that was
B. splendens and B. decadactylus in the Azores Archipelago based on obtained from the database of the Department of Oceanography and
all available data to test the hypothesis that the stocks of these spe- Fisheries, University of the Azores (DOP/UAç), collected during the
cies in the ICES Subarea 10a2 represent a FMU. period of 1990–2016 as part of the DCF. Sampling was designed to
cover the main ports of the Azores and included completing standar-
dised fishing questionnaires (n = 9282) with the captains of the bot-
2 | MATERIALS AND METHODS
tom longline vessels during the landings. Each record report included:
the vessel identification, the dates of departure and return to the port
2.1 | Study area and detailed information on fishing operations, including the number
The Azores Archipelago is in the triple junction of the North American, of hooks per set, number of sets per trip, gear characteristics, fishing
Eurasian and African tectonic plates and represents the only coastal area and total mass of catch and catch by size categories for each spe-
margins associated with the Mid-Atlantic Ridge on the North Atlantic cies landed.
SANTOS ET AL.
FISH 3

40°

Corvo

Flores Serreta
Bank
Graciosa
39°
Terceira
Faial São Jorge
Condor
Bank

Acores Pico
38° Bank São Miguel.
Princesa Alice
Bank
Formigas
Voador Bank

Monte Alto
37°
Santa Maria

31° 30° 29° 28° 27° 26° 25°

FIGURE 1 The Azores archipelago (NE Atlantic Ocean) with the location of the nine islands (Corvo, Flores, Faial, Pico, São Jorge, Graciosa,
Terceira, São Miguel and Santa Maria) and main seamounts

2.3 | Data analysis 800 m, a mode at 400–450 m (Figure 2) and most of the abundance
(89%) at 100–600 m strata (Figure 2). The LF distribution ranged from
The catch per hook value (i.e., catch per unit effort; CPUE) was calcu-
15 to 40 cm with a mode at c. 27 cm (Figure 3). Similar lLF composi-
lated for each species, surveyed area and station stratum and an index
tion was observed by area (islands and seamounts) but with about
of relative abundance in number (or mass) was obtained by multiplying
70% of the total abundance found on the island regions (Figure 3).
each of these CPUE values by the corresponding area size. The average
Larger individuals were found at the deeper strata (Figure 4). Annual
relative abundance value for each area and stratum was then calcu-
mean LF presented a recovery period from 2008 to 2013 after
lated. The annual abundance values for each area and for the Azores
13 years of decline (Figure 5). However, the mean LF has been declin-
were computed by summing the abundance values across strata and
ing again since 2016 (Figure 5). The LF composition of commercial
across areas, respectively. Finally, the bootstrap method (Efron & Tib-
landings varied between 12 and 53 cm with a mode of c. 30 cm
shirani, 1986) was applied to calculate confidence intervals and check
(Figure 6). No information about LF by area or depth is available or is
statistical differences for the annual estimated abundance values.
reported in the DCF dataset.
Relative indices of abundance for the Beryx spp. caught by Azor-
This species presented a high and significant (P < 0.05) inter-
ean commercial longline fleet were estimated by generalized linear
annual variability in the various abundance indices during the sur-
modelling approach using a hurdle (Δ) model (Lo et al., 1992; Ortiz &
veyed period; the survey index showed a decreasing trend until 2000,
Arocha, 2004; Zuur & Ieno, 2016). The standardisation protocols
low and stable trend during the period 2000–2008, some recovery
assumed a hurdle model (zero-altered log normal) with a binomial
from 2010 to 2013 and again a decrease afterwards (Figure 7). By
error distribution and logit link function for modelling the probability
contrast, analysis of the commercial landings index showed an oscilla-
that a null or positive observation occurs (proportion of positive
tion over time, with a general decreasing trend from the year 2009
catches) and a log-normal error distribution with an identity link func-
(Figure 7).
tion for modelling the positive catch rates on successful trips.
All the analyses were conducted in R 3.4.3 (www.r-project.org)
with additional packages lmtest (Zeileis & Hothorn, 2002), lattice 3.2 | Beryx decadactylus
(Sarkar, 2008), influence.ME (Nieuwenhuis et al., 2012), HLMdiag
Beryx decadactylus was caught in all statistical areas sampled within
(Loy & Hofmann, 2014), lme4 (Bates et al., 2015) and lsmeans
the Azores region with a typical depth distribution between 200 and
(Lenth, 2016).
950 m and a peak in abundance between 700–750 m (Figure 2). This
resource seemed to be equally distributed along the depth strata, with
about 49% of the total abundance found at 200–600 m and 51% on
3 | RESULTS
the deepest (600–950 m) ones (Figure 2).
The LF distribution of this species ranged from 15 to 67 cm with a
3.1 | Beryx splendens mode at c. 42 cm, but with a spread distribution (Figure 3). Similar LF
Beryx splendens was caught in all statistical areas sampled within the compositions were found by area (islands and seamounts) but with about
Azores region with a typical depth distribution between 100 and 60% of the total abundance found on the seamounts regions (Figure 3).
4
FISH SANTOS ET AL.

(a) (a)
14 55

12 50

10
45
8
40
6
35
4
Relave abundance index

2 30

0 25

(b) 20

LF (cm)
14
12 (b) 55

10 50
8
45
6
40
4
35
2
0 30
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0– 0
0
15 15
20 25
25 25
30 30
35 35
40 40
45 45
50 50
55 55
60 60
65 65
70 70
75 75
80 80
85 85
90 90
95

25
0–
10

Depth strata (m) 20

200–250

250–300

300–350

350–400

400–450

450–500

500–550

550–600

600–650

650–700

700–750

750–800

800–850

850–900
FIGURE 2 Mean (± 95% CI; –) abundance index by depth strata of
(a) Beryx splendens and (b) B. decadactylus 1995–2017 from the annual
Azorean spring bottom longline survey Depth strata (m)

FIGURE 4 Mean (± 95% CI) fork length (LF) by depth strata of


(a) Beryx splendens and (b) B. decadactylus 1995–2017 from the annual
Azorean spring bottom longline survey

(a) 3.5

3.0 (a) 45

2.5
40
2.0
35
1.5

1.0 30
Relave abundance index

0.5 25
0
20
LF (cm)

(b) 3.5
(b) 45
3.0
40
2.5

2.0
35

1.5 30

1.0
25
0.5
20
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017

0
15 18 21 24 27 30 33 36 39 42 45 48 51 54 57 60 63 66
LF (cm) Year

FIGURE 3 Fork length (LF)-frequency (relative abundance) FIGURE 5 Annual mean (± 95% CI) fork length (LF) of (a) Beryx
distribution by area of (a) Beryx splendens and (b) B. decadactylus splendens and (b) B. decadactylus, 1995–2017 from the Azorean spring
1995–2017 from the annual Azorean spring bottom longline survey. bottom longline survey. n.b. There is no data for the years 1998,
( ) Islands and ( ) Seamounts 2006, 2009, 2014 and 2015
SANTOS ET AL.
FISH 5

(a) 25000 (a) 3.5


3
20000
2.5

15000 2

1.5
10000
1

CPUE (kg 10–3 hooks)


0.5
Landings in number

5000
0
0
(b) 3.5
(b) 3500 3
3000 2.5
2500
2
2000
1.5
1500
1
1000
0.5
500
0
0

1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
9 13 17 21 25 29 33 37 41 45 49 53 57 61 65
LF (cm) Year

FIGURE 6 Fork length (LF)-frequency distribution of (a) Beryx FIGURE 7 Nominal ( ) and standardised ( ) annual catch per unit
splendens and (b) B. decadactylus 1990–2016 from the landings of the effort (CPUE) series of (a) Beryx splendens and (b) B. decadactylus
Azorean commercial bottom longline fleet 1990–2017 from landings of the Azorean commercial bottom longline
fleet and annual CPUE series from long-line surveys ( ). CPUE
Larger individuals were found in the deeper strata (Figure 4) and annual series were scaled by normalizing relatively to the time series mean
mean LF presented a great variation over time (Figure 5). The LF distribu-
tion of commercial landings varied between 12 and 53 cm with a mode hypothesize that B. splendens comprises a single population that
at c. 36 cm (Figure 6). No information about LF by area or depth was migrates between the Corner Rise and the Azores banks (Alekseev
available or was reported in the DCF dataset. et al., 1987). This hypothesis, however, is largely based on non-
This species also presented a high and significant (P < 0.05) inter- representative length–age composition data of B. splendens that sug-
annual variability in the abundance index during the surveyed period gest only small immature fish inhabit the Azores whilest the Corner
(Figure 7). With regard to the survey-derived index of abundance, a Rise is inhabited mostly by large mature fish (González-Costas, 2015).
peak of abundance was observed in 1995 that fell to low level in However, this species does spawn around the Azores and all sizes and
1997 and then levelled off until 2003. It then increased until 2007, phases of the life cycle occur on the fishing and sampled survey area
had a slight decreasing trend during the following 3 years, showed (ICES, 2017; Rico et al., 2001) and the same occurs in the Corner Rise
some recovery in 2011, but soon fell again (Figure 7). From the com- (Vinnichenko, 1997). Furthermore, there is no single record on mature
mercial landings analyses, the index trends showed a peak in 1991, B. splendens migrations in the open ocean (Kotlyar, 1996). In general,
followed by a decreasing trend until 1997 and less variation thereafter this species reaches the first sexual maturation around 20–35 cm
(Figure 7). (González et al., 2003; Kozlov, 2014; Lehodey et al., 1997), but the
species is remarkably plastic in this regard and changes according to
the area in response to the different exploitation levels (González
4 | DISCUSSION
et al., 2003). However, this discrepancy in size at first sexual matura-
tion is also due to the unknown stock structure for the Atlantic Ocean
Little is known about the B. splendens population structure. Hoarau &
Borsa (2000) have suggested genetic homogeneity between Pacific resulting in sampling bias for life-history parameter estimation (FAO,

and Atlantic populations but they included only four specimens from 2016; Friess & Sedberry, 2011a,b). Within the Azores region, different

the Atlantic Ocean and more work is needed to test this gene flow sizes at first sexual maturation were found: 23 cm (González et al.,
idea (Friess & Sedberry, 2011a). On a more local scale, genetic differ- 2003), 24.7 cm (Isidro, 1996) and 35.5 cm (Pereira & Pinho, 2012).
entiation was observed among Macaronesian populations from the Based on these results, practically all fish caught during the studied
Canary Islands, Madeira and the Azores suggesting a role for oceanic period in the commercial fishery and in the surveys were mature
currents, as geographical barriers among the Macaronesian archipela- individuals.
gos preserving distinctive haplotypes (Schönhuth et al., 2005). Beryx splendens in spawning condition (Isidro, 1996) and size
Beryx splendens is found also on seamounts along the Mid-Atlantic (23–35 cm) are observed around the Azorean seamounts and islands.
Ridge, north and south of the Azores. In this area, some scientists This is an indication that reproduction in this species may not involve
6
FISH SANTOS ET AL.

large migrations and that B. splendens reproduce in the regions they Beryx decadactylus has a worldwide distribution and sometimes
inhabit as suggested by authors Masuzawa et al. (1975), Alekseev taken in the same fishery as B. splendens (FAO, 2016). Most of the
et al. (1986) and Lehodey et al. (1997). The variation in the LF – population genetics studies of Beryx species focus on the economi-
frequency distribution of this species populations as a function of cally more important B. splendens. However, for B. decadactylus a
depth was also found in other studies (Alekseev et al., 1986; Lehodey recent study comparing the mtcr gene sequences of fish caught on
et al., 1997; Massey & Horn, 1990). Alekseev et al. (1986) and Leho- the western North Atlantic Ocean (off the south-eastern USA) and
dey et al. (1997) suggested that the vegetative zones, in which juve- Azores areas suggested no differences between these populations
niles of B. splendens grow until they reach maturity, are seamounts (Friess & Sedberry, 2011a,b). On the other hand, a previous study
with shallow depths (c. 390 m) and reproductive zones inhabited by showed some evidence for localized genetically different populations
mature individuals are waters with depths between 500 and 750 m between Azores and Cape Verde regions on the eastern North Atlan-

and that the early life stages would be carried from the reproductive tic Ocean (Aboim, 2005).

zone to the vegetative zone by currents. A similar pattern could be Beryx decadactylus is considered to have a longevity of 69 years

applied to the B. splendens population of the Azores. The combined off the south-eastern USA (Friess & Sedberry, 2011b), while in the
Azores they were aged up to 10 years, 11 years in Madeira and
results from this and previous studies show that the spawning proba-
9 years in the Canary Islands (Krug et al., 2010). The size of the first
bly occurs in area inhabited by adults (600–900 m), eggs and larvae
sexual maturation occurs 27–30 cm for males and 30–33 cm for
develop in surface-water layer (Mundy, 1990), juveniles have a pelagic
females (Estácio et al., 2001; Isidro, 1996). However, although these
stage (Isidro, 1996) and grow in demersal zones that are shallower
sizes have been recorded in the fishing and survey data, practically no
(< 600 m) than areas inhabited by adults (Figure 8). Nevertheless, fur-
sexually mature individuals were caught on the Azores area and aggre-
ther studies are required to confirm this scheme of its life cycle.
gations were not reported to date (ICES, 2010; Isidro, 1996). Larger
Beryx splendens is also found on seamounts along the Mid-Atlantic
and mature individuals were found in areas off the south-eastern USA
Ridge, north and south of the Azores, being deeply exploited by the
(Friess & Sedberry, 2011b). These results seem to suggest a migratory
Soviet–Russia fleets, particularly to the north, since the 70s
dynamic of adults from western areas like the one observed for other
(Vinnichenko, 2002). Little is known about movements of this species
deep-water species (Friess & Sedberry, 2011b). Nevertheless, little is
between the Azores EEZ area and other seamounts on the Mid-
known about movements of B. decadactylus between the Azores EEZ
Atlantic Ridge. However, it is proposed that the Mid-Atlantic Ridge
and other areas and how the LF -frequency distribution varies
seamounts can support independent management units, since no
between fishing grounds and different depths. In the present study,
recovery was detected after the seamounts depletion by the
the results indicated that this species is more abundant in the sea-
Soviet–Russia fleets (see Vinnichenko, 2002), even knowing about the
mounts areas, with a bigger-deeper trend but further studies consider-
high mobility of this species (Silva and Menezes, 1996).
ing an entire population are necessary to investigate its life cycle
(Figure 8).
Beryx splendens The results from this study, combined with previous ones of
Beryx decadactylus B. splendens and B. decadactylus indicate that the stocks of these spe-
3 2 cies in the Azores (ICES Subarea 10a2) may be a FMU only for
B. splendens, while B. decadactylus is a single stock for the North
Atlantic Ocean. Detailed information is available on the Azores for
stock assessment of splendid alfonsino in the Azores. So we recom-
4
mend that exploratory analysis for analytical assessment be performed
1 considering this resource as a local management unit.

ACKNOWLEDGEMENTS
6
7
Ricardo Medeiros (ImagDOP/UAz) is thanked for generation of the
5 ?
map. The authors thank all who participated in field surveys and sam-
pling processing onboard the R.V. Arquipélago. This study resulted
FIGURE 8 Diagram of life cycle of Beryx spp. around Azores islands from a recommendation of the International Council for the Explora-
and seamounts and connectivity hypotheses among different habitats. tion of the Sea (ICES) during the Working Group on the Biology and
Beryx splendens: (1) spawning occurs in area inhabited by adults, and
Assessment of Deep-sea Fisheries Resources (WGDEEP) in 2018.
(2) eggs and larvae develop in surface-water layer; (3) juveniles have a
pelagic stage and grow in demersal zones that (4) are shallower than
areas inhabited by (5) adults. B. decadactylus: (6) more abundant in ORCID
seamount areas but no sexually mature individuals were caught on Régis V. S. Santos https://orcid.org/0000-0002-4167-3573
the Azores exclusive economic zone (EEZ) and aggregations were not
Ana M. Novoa-Pabon https://orcid.org/0000-0002-7194-7176
reported up to date; (7) larger and mature individuals seem to migrate
to the western areas off the Azores but little is known about this Hélder M. Silva https://orcid.org/0000-0001-7181-1976
behaviour Mário R. Pinho https://orcid.org/0000-0002-8045-2546
SANTOS ET AL.
FISH 7

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deep-sea demersal fishes from the Azores—North Atlantic. (Master’s the-
and ecology of a deep-demersal fish, alfonsino Beryx splendens, over
sis). University Southampton, Hampshire, England, 167 p.
the seamounts off New Caledonia. Marine Biology, 128, 17–27.
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roscale water circulation, ontogenetic geographical differentiation and
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1986/C:10. Hydrographic Committee. 16 p.
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