Exploring composition and behaviour of fish fauna by in situ observations in the Bari Canyon...

ORIGINAL ARTICLE

Exploring composition and behaviour of fish fauna byin situ observations in the Bari Canyon (Southern AdriaticSea, Central Mediterranean)Gianfranco D’Onghia, Francesca Capezzuto, Angela Carluccio, Roberto Carlucci, Agnese Giove,Francesco Mastrototaro, Michele Panza, Letizia Sion, Angelo Tursi & Porzia Maiorano

Department of Biology and CoNISMa LRU, University of Bari Aldo Moro, Bari,Italy

Keywords

Behaviour; canyon; feeding habit; fish fauna;

Mediterranean.

Correspondence

Gianfranco D’Onghia, Department of

Biology, University of Bari Aldo Moro – Via E.

Orabona 4, 70125, Bari, Italy.

E-mail: [email protected]

Accepted: 8 February 2014

doi: 10.1111/maec.12162

Abstract

Canyons play a fundamental role in enhancing the abundance and diversity of

marine organisms through the transport of organic matter and food resources,

the presence of complex physical habitats and the absence of trawl fishing.

During four baited lander deployments carried out in the Bari Canyon (South-

ern Adriatic Sea, Central Mediterranean), at depths of 443–788 m, about 43 h

of video records were taken, for a total of 619,200 video frames. A total of 12

benthopelagic fish species (five chondrichthyes and seven osteichthyes) were

identified. The blackspot seabream (Pagellus bogaraveo) was the most often

observed fish species. The depth of 787 m represents a new depth record for

this fish in the Adriatic Sea. Groups of up to 40 individuals of P. bogaraveo

were attracted to the bait and were shown in single frames. The individuals

were observed both exploring the area and feeding actively on the bait. The

European conger (Conger conger) was recorded at each deployment. Clear

scavenger behaviour was also observed in this teleost fish and in the shark

Etmopterus spinax. The shark species Centrophorus granulosus and Hexanchus

griseus, which are considered ‘vulnerable’ on the published IUCN Mediterra-

nean Regional Red List, were also recorded but, although attracted by the bait,

they were never seen feeding on it. Other fish species, harvested on fishing

grounds, such as Merluccius merluccius, Helicolenus dactylopterus and Polyprion

americanus, were also recorded. This study represents the first in situ documen-

tation, at very low impact, of the fish fauna in the Bari Canyon, providing new

insights into its small scale distribution and behaviour, the first in situ direct

observation of the variable feeding behaviour of P. bogaraveo and its gregarious

habits, as well as indicating that this canyon could act as a refuge area for

species that are vulnerable to fishing on the open slope.

Introduction

Submarine canyons are complex structures capable of

enhancing the heterogeneity of the slope and therefore

play an important role in the functioning of the deep sea

ecosystem (Danovaro et al. 2010; Wurtz 2012). In fact,

canyons act as trapping areas for sediments and provide a

direct pathway for sediment transport and dense water

cascading from the continental shelf to the deep basin,

enhancing the transport and availability of particles and

organic matter (Turchetto et al. 2007; Tesi et al. 2008;

Canals et al. 2009). The effect of canyons on local circula-

tion can enhance the abundance and the diversity of mar-

ine organisms, through the transport of sediment and

phytodetritus and the presence of complex physical habi-

tats (Vetter & Dayton 1998; Ramirez-Llodra et al. 2010).

Higher macro- and megafauna biomass as well as spawn-

ing and recruitment sites have been detected in the Medi-

terranean canyons (e.g. Company et al. 2012; Wurtz

2012; Tecchio et al. 2013 and references therein).

Marine Ecology (2014) 1–16 ª 2014 Blackwell Verlag GmbH 1

Marine Ecology. ISSN 0173-9565

Moreover, canyons can be considered as biodiversity hot-

spots with high megafauna diversity and high rates of

endemism (Danovaro et al. 2010 and references therein;

Ramirez-Llodra et al. 2010 and references therein).

The enhancement of trophic resources, accelerated cur-

rents and structural heterogeneity also make these canyons

favourable habitats for suspension feeders such as cold-

water corals (CWC) (Freiwald et al. 2009; Orejas et al.

2009). CWC have also been recorded in the Bari Canyon

System (Southern Adriatic Sea, Central Mediterranean)

using a remotely operated vehicle (ROV) (Freiwald et al.

2009). This canyon has been investigated on geomorpho-

logical, oceanographic and sedimentological aspects (e.g.

Ridente et al. 2007; Trincardi et al. 2007; Turchetto et al.

2007; Tesi et al. 2008; Rubino et al. 2010; Carniel et al.

2012). Information on the distribution and abundance of

megafauna in the Bari Canyon is mainly related to benthic

sessile species (Freiwald et al. 2009; Bo et al. 2012; Sanfi-

lippo et al. 2013; Angeletti et al. 2014) while knowledge of

the fish fauna is limited to the recording of four teleost

fishes using ROV (Angeletti et al. 2014). This lack of

knowledge can be explained by the inaccessibility of the

area to traditional sampling gear and trawl nets, making

the sampling of mobile megafauna quite difficult. Longline

fishing only occurs on the margins of the Bari Canyon,

mostly during summer. The walls and sea floor of the can-

yon are very rough three-dimensional structures that form

habitats, such as hard ground, boulders and coral branches,

which can tear or entangle longlines.

Video inspections are generally less invasive for fragile

deep-sea ecosystems and can provide information on

small scale species distribution, activity and behaviour

(e.g. Bailey & Priede 2002; Uiblein et al. 2003; Costello

et al. 2005 and references therein; Ross & Quattrini 2007;

King et al. 2006, 2008 and references therein; Capezzuto

et al. 2012). However, several difficulties and limitations

should be taken into consideration during the analysis of

fish from deep sea environments. ROV sampling shows

limitations, among which are those mainly related to

noise and light created by the ROV that may either scare

away or attract fish, thus causing under- or overesti-

mations of the true abundances (Trenkel et al. 2004;

Costello et al. 2005; Lorance & Trenkel 2006; Stoner et al.

2008; Ryer et al. 2009). Such underestimations of species

that avoid the ROV and over estimates of species that are

attracted by the presence of the ROV are problems that

should be considered, in particular for species which are

rarely seen. Benthic landers produce less disturbance in

gathering similar data but are vulnerable to their own

disadvantages. Landers will only attract scavenging fish

and so are highly selective. The time required to explore

unknown deep-sea bottoms suitable for setting up and

precisely positioning the equipment, as well as the time

required at a single station between deployment and

recovery and the many assumptions made in abundance

estimates are other sampling limitations involved in the

use of landers (Priede & Merrett 1996; Roberts et al.

2005; Jamieson et al. 2006; Bailey et al. 2007; Cousins

et al. 2013).

During a survey carried out as part of the RITMARE

Italian project aimed at studying biodiversity and the rela-

tive ecosystem services in the Bari Canyon, data on the fish

fauna were collected using a baited lander equipped with

two digital cameras. In this work, the video records were

used: (i) to explore the composition of the benthopelagic

fauna in this canyon; (ii) to investigate their small-scale

distribution and activity; and (iii) to examine the behav-

iour of the individuals by species. Due to the difficulties

involved in the study of mobile bentopelagic fauna in a

habitat with complex and rugged bathymetry such as the

Bari Canyon, the work is explorative in character.

Material and Methods

Study area

The Bari Canyon System (BCS) is a complex morphologi-

cal structure that breaches the Southwestern Adriatic shelf

with a west–east trend. It is 10 km wide and 30 km long

and is between 200 and 1700 m in depth (Ridente et al.

2007; Trincardi et al. 2007). The BCS is characterized by

two main branches separated by a mounded relief, a

slope-confined trough between 500 and 700 m in depth.

The southern branch is wider (about 4 km), markedly

asymmetric and U-shaped with a very steep wall (Ridente

et al. 2007; Trincardi et al. 2007).

As part of the RITMARE project funded by the Italian

Ministry of University and Research, the MEMO (Marine

Environment MOnitoring system) lander was deployed in

the southern branch of the Bari Canyon (Southern Adri-

atic Sea, Central Mediterranean) in May–June 2012 using

the R/V MINERVA UNO. The four deployments are pre-

sented in Fig. 1 with indications of the deployment

depths.

Baited lander deployments

The MEMO lander, developed as part of the CoralFISH

EU_7FP project, consists of a stainless steel metal frame,

two digital cameras (Sony ICX414) with two LED lights

(12 V, 700 mA, 360 lumen), an electronic compass, incli-

nometer and altimeter, a multiparametric probe, a dopp-

ler current meter, four deep-sea batteries for power

supply, and an ICT infrastructure capable of managing

the entire system (Fig. 2). On the sea bed the lander is

linked by a zinc-coated steel cable to buoys which keep

2 Marine Ecology (2014) 1–16 ª 2014 Blackwell Verlag GmbH

Fish fauna by in situ observations D’Onghia, Capezzuto, Carluccio, Carlucci, Giove, Mastrototaro, Panza, Sion, Tursi & Maiorano

the cable under tension and then to a surface floating sig-

nalling buoy. Continuous connection is maintained via

an acoustic modem with an onboard PC software plat-

form, making images and sensor data available.

A total of four deployments between 443 and 788 m

were carried out, for a total time of about 43 h of video

recordings (Table 1). As the MEMO system takes four

video frames each second, a total of 619,200 video frames

were taken by each camera. During each deployment the

lander was baited with eight specimens of Scomber scom-

brus (~700 g). The bait produces an odour plume stretch-

ing downstream from the lander and attracting animals

in view of the camera (Bailey et al. 2007). The hydro-

graphic parameters at each station were also recorded by

means of CTD and a current meter. The mean salinity

recorded on the bottom was 38.72 psu, the mean temper-

ature values varied from 13.32 to 13.38 °C, and those of

the current speed from 0.06 to 0.14 m�s�1.

Data collection and analysis

The different species were identified by their morphologi-

cal characteristics observed in the video frames. In

addition, with the exception of Pagellus bogaraveo, the

single individuals of all recorded species could often be

distinguished by their different sizes and morphological

features, such as scars on the skin. Thus, with the excep-

tion of P. bogaraveo, a total number by species was

recorded for each deployment. According to King et al.

(2006, 2008), the arrival time of the first individual (1st

arrival time) and maximum number of individuals

observed at the bait per deployment (MaxN) were

recorded. Since fish numbers gradually increased after

lander touchdown but with individuals moving in and

out of camera view, the number of fish visible at any

given time during a sequence varied. Thus, the percentage

of frames with individuals was recorded in each deploy-

ment for each species.

As significant numbers of P. bogaraveo individuals were

observed, the variations in the visual count detections

between day and night hours and current velocity were

examined following Aguzzi et al. (2003, 2006) and Doya

et al. (2014). Thus, the maximum numbers of individuals

in the field of view were recorded every 30 s (MaxN30s).

The mean of MaxN30s for each hour of deployment, with

standard deviation, was computed to obtain a mean

17°4'0'' E 17°8'0'' E 17°12'0'' E 17°16'0'' E 17°20'0'' E 17°24'0'' E 17°28'0'' E

17°4'0'' E 17°8'0'' E 17°12'0'' E 17°16'0'' E 17°20'0'' E 17°24'0'' E 17°28'0'' E

17°32'0'' E

17°32'0'' E

41°2

4'0'

' N41

°20'

0'' N

41°1

6'0'

' N41

°12'

0'' N

41°2

4'0'

' N41

°20'

0'' N

41°1

6'0'

' N41

°12'

0'' N

Fig. 1. Study area: location within the Mediterranean of the Bari Canyon and bathymetric map with indication of the MEMO lander deployments.


D’Onghia, Capezzuto, Carluccio, Carlucci, Giove, Mastrototaro, Panza, Sion, Tursi & Maiorano Fish fauna by in situ observations

fluctuation plot. Phase duration and timing was assessed

using the MESOR (midline estimating statistic of rhythm)

estimated by re-averaging all hourly values and represent-

ing the resulting mean as a horizontal threshold line on

the fluctuation plots (Aguzzi et al. 2003, 2006; Doya et al.

2014). The fluctuation of the hourly mean current veloc-

ity (m�s�1) during the deployment was superimposed on

the fluctuation plot of the visual fish count.

Sizes of fishes were measured from images using

IMAGE J 1.46q, a Java-based public domain program

developed at the USA National Institutes of Health, Beth-

esda, MD, USA (available at, http://rsb.info.nih.gov/ij/

index.html). The software was calibrated using the

within-image reference scale (bait plate, 25 cm diameter).

Fishes were only measured if the full length of the body

was visible at the level of the reference scale. The size was

expressed as total length (TL, cm). To avoid re-measuring

the same individual, the fishes were only measured at

peak abundance in the field of view (MaxN) in the differ-

ent deployments. However, specimens of most species

were often distinguished in relation to their different

sizes, colour and scars on the skin. For P. bogaraveo the

length–frequency distributions were also computed at the

depths where individuals with significant numbers were

detected and measured.

Position with respect to the bottom, reaction to the

lander and activity of the individual organisms were clas-

sified in terms of ‘Fauna Position’, ‘Fauna Reaction’ and

‘Fauna Activity’, respectively, as described in Uiblein

et al. (2003), Trenkel et al. (2004) and Costello et al.

(2005). In particular, Fauna Position was recorded in

three categories: water column (1–2 m above the

Fig. 2. The CoNISMa MEMO lander equipment with indication of the different components.

Table 1. Data related to the MEMO lander deployments in the Bari Canyon.

Date Station Latitude Longitude Depth (m)

Salinity

(psu) T(°C)

Mean

current (m/s)

Start

video

End

video

Time of

video record

(h, m)

29/5/2012 St. 1 41°18.40850 N 17°09, 06220 E 443 38.72 13.37 0.12 11:23 18:37 7.14

30/5/2012 St. 2 41°17.70930 N 17°17,49490 E 787 38.72 13.32 0.06 09:58 17:37 7.39

30–31/05/2012 St. 3 41°17.71270 N 17°17,71260 E 788 38.72 13.34 0.14 18:50 07:35 12.45

31/05–1/06/2012 St. 4 41°14.48760 N 17°20,34150 E 577 38.72 13.38 0.06 11:35 04:01 16.26



bottom); near the sea bed (within 0.5–1 m); and on the

sea bed. Fauna Reaction was classified in three categories:

no reaction; avoidance-disturbed; and attraction. Fauna

Activity was divided into seven categories: actively swim-

ming; quietly swimming; hovering; moving on the sea

bed; resting on the sea bed; feeding; and feeding on bait.

Results

Species composition

A total of 12 benthopelagic fish species (five chondrich-

thyes and seven osteichthyes) were identified (Table 2,

Fig. 3). The highest number of species (8) was observed

at the shallowest station (443 m). Most species were

recorded during daylight hours. Only one individual was

observed during one deployment for the species Cen-

trophorus granulosus, Galeus melastomus, Hoplostethus

mediterraneus and Polyprion americanus.

Pagellus bogaraveo was the fish most frequently

observed at the two shallowest stations: in 85.16% of the

video frames at 443 m and 51.98% at 577 m. One indi-

vidual of this fish was observed at 787 m, which is a new

depth record of this fish in the Adriatic Sea. Its total

number per deployment is not reported in the Table 2

due to the impossibility of distinguishing individuals.

First arrival time for P. bogaraveo was 9 min at 443 m

with a maximum number of 40 individuals in the frames

and 90 min at 577 m with a maximum number of 18

individuals in the frames (Fig. 4). The individuals mea-

sured at 443 m had sizes between 12.5 and 37.5 cm TL

with a mean length of 21 � 5.1 cm TL; those measured

at 577 m ranged from 12.5 to 39 cm TL and the mean

length was 25 � 8.7 cm TL (Fig. 5).

A significant increase in the visual counts of P. bogaraveo

was detected during the afternoon at 443 m and near sun-

set at 577 m. Superimposing the current velocity on the

fluctuation in the visual fish count seems to indicate that

animal detections increase significantly when water speed

decreases at both stations (Fig. 6). Water speed at 577 m,

when day and night hours were investigated, showed a

diurnal oscillation with a higher peak at night and a smaller

one in daytime.

Other fishes frequently observed at the shallowest

deployment were the teleosts Helicolenus dactylopterus

(25.98%) and Conger conger (9.97%). Both species were

recorded during day and night hours. The latter was the

only fish recorded in all four deployments. The maxi-

mum number (7) was observed at 788 m, where it

arrived sooner (12 min) than at the other stations. The

greatest sizes were shown at the greatest depths. The

other teleosts and the cartilagineous fishes were observed

less frequently. The two shark species G. melastomus

(49 cm TL) and Etmopterus spinax (33 and 45 cm TL)

arrived at the bait in a comparable time, 28 min at

443 m and 26 min at 787 m, respectively.

Large individuals (61 and 62 cm TL) of the commer-

cial target species Merluccius merluccius were recorded at

the two shallowest stations during daytime.

Species behaviour

The list of the species recorded with indication of type of

behaviour (position, reaction and activity) is presented in

Table 3. All the species were observed swimming near the

sea bed. However, Pagellus bogaraveo was mainly

recorded swimming, both actively and quietly, in the

water column. It was often observed hovering in front of

the lander near the bait. Galeus melastomus, Conger con-

ger, Helicolenus dactylopetrus and Merluccius merluccius

were also recorded on the sea bed. None of the observed

species seemed to be clearly disturbed by the lander. Indi-

viduals of most species either showed no reaction or were

attracted to the lander. Etmopterus spinax, C. conger and

P. bogaraveo were observed investigating, attacking and

ingesting the bait (Fig. 7). Individuals of P. bogaraveo

were mostly observed in groups swimming in the water

column and near the sea bed attracted to the baited

lander. Members of the group were observed forming

irregular group shapes. The groups appeared to be rather

disorganised near the lander, with most individuals

actively swimming and investigating, others quietly swim-

ming in a circle, and others hovering. They sometimes

showed no reaction. Individuals in the groups were some-

times observed swimming together in a coordinated fash-

ion (Fig. 8). Fish were mostly observed staying on the

bait for a long time, accumulating in numbers within the

camera’s view.

The first specimens ahead of the group often attacked

the bait, removing flesh and even single fish from the bait

plate (Fig. 9). The other individuals generally continued

to attack the same fish primarily utilized on the bait

plate, exhibiting a feeding frenzy. Some individuals were

observed ingesting the bait and others also feeding on the

fast-moving plankton in the water column and near the

bottom. The fish often remained within the camera’s view

even after all the bait had been totally consumed.

Helicolenus dactylopterus was also observed feeding on

the plankton and small crustaceans moving or resting on

the sea bed. It was never observed to feed on the bait.

Lepidopus caudatus was shown feeding in the sediment.

Apart from E. spinax, the other shark species were only

observed being attracted by the bait and investigating it

but without feeding. The wreckfish P. americanus was

observed quietly swimming near the sea bed with no

reaction to the lander.



Table 2. First arrival time on bait, total number of individuals observed per deployment (Total N), maximum number of individuals observed at

the bait per deployment (Max N), percentage of frames in which each species was visible, total length (TL) and presence during day or night

hours (Day-Night). (n.m. = not measured)

Species

Deployment 1 2 3 4

Depth (m) 443 787 788 577

Chondrichthyes

Centrophorus granulosus First arrival time (min) 116

Tot N 1

Max N 1

% of frames 1.08

TL (cm) 89

Day-Night D

Dalatias licha First arrival time (min) 129

Tot N 2

Max N 1

% of frames 1.44

TL (cm) 96

Day-Night D

Etmopterus spinax First arrival time (min) 26

Tot N 2

Max N 1

% of frames 0.84

TL (cm) 33–45

Day-Night D

Galeus melastomus First arrival time (min) 28

Tot N 1

Max N 1

% of frames 0.57

TL (cm) 49

Day-Night D

Hexanchus griseus First arrival time (min) 268 219

Tot N 1 1

Max N 1 1

% of frames 0.04 0.12

TL (cm) n.m. n.m.

Day-Night D D

Osteichthyes

Conger conger First arrival time (min) 44 47 12 174

Tot N 5 3 7 2

Max N 2 1 2 1

% of frames 9.97 2.74 5.98 1.4

TL (cm) 66 � 17 106 � 20 117 � 12 67–110

Day-Night D D D-N D-N

Helicolenus dactylopterus First arrival time (min) 305 57 76

Tot N 2 2 2

Max N 2 1 1

% of frames 25.98 5.47 3.20

TL (cm) 16–25 22–29 25–29

Day-Night D D D-N

Hoplostethus mediterraneus First arrival time (min) 2

Tot N 1

Max N 1

% of frames 0.40

TL (cm) 19

Day-Night D



Discussion

The present results represent the first in situ documenta-

tion on the fish fauna composition and behaviour in the

Bari Canyon. The fish fauna distributed in this canyon is

not taxonomically different from the overall regional

fauna. Apart from Pagellus bogaraveo, which was shown

at the depth record of 787 m for Adriatic Sea, the vertical

distribution observed in the species falls within the depth

range known for each of them (Ungaro et al. 1998a,b;

Relini et al. 2000, 2010; Vacchi & Serena 2010). The pres-

ence of Helicolenus dactylopterus and P. bogaraveo in the

Table 2. Continued

Species

Deployment 1 2 3 4

Depth (m) 443 787 788 577

Lepidopus caudatus First arrival time (min) 408 259

Tot N 1 2

Max N 1 1

% of frames 0.003 0.03

TL (cm) n.m. n.m.

Day-Night D D

Merluccius merluccius First arrival time (min) 107 362

Tot N 1 1

Max N 1 1

% of frames 0.66 0.41

TL (cm) 61 62

Day-Night D D

Pagellus bogaraveo First arrival time (min) 9 214 90

Tot N – – –

Max N 40 1 18

% of frames 85.16 0.11 51.98

TL (cm) 21 � 5.1 25 � 8.7

Day-Night D D D-N

Polyprion americanus First arrival time (min) 268

Tot N 1

Max N 1

% of frames 0.13

TL (cm) n.m.

Day-Night D

Fig. 3. Images of most benthopelagic fish species recorded by MEMO lander in the Bari Canyon.



Bari Canyon was previously recorded at depths between

382 and 479 m during a recent ROV exploration (Ange-

letti et al. 2014).

Almost all the fish species found have been already

recorded in other Mediterranean canyons (e.g. Stefanescu

et al. 1994; Ramirez-Llodra et al. 2010; Company et al.

2012; Farrugio 2012; Watremez 2012; Tecchio et al.

2013). Even though there is no indication of a specialist

canyon fauna, some species, such as Centrophorus granu-

losus, Hexanchus griseus and Polyprion americanus, which

consist of rare populations and are less accessible to fish-

ing gear, are reported in the studies carried out in the

canyons using underwater video systems (e.g. Watremez

2012) but generally not collected using trawl nets in these

geomorphological structures (e.g. Stefanescu et al. 1994;

Ramirez-Llodra et al. 2010; Company et al. 2012).

Considerable information is available on the abundance

of the demersal ichthyofauna on the trawlable bottoms of

the Southern Adriatic Sea at depths shallower than 800 m

(e.g. Ungaro et al. 1998a,b and references therein; Relini

et al. 2000, 2010 and references therein). Large specimens

of the species observed in this study are generally caught

A B

Fig. 4. Maximum number of individuals (N = 40; N = 18) of Pagellus bogaraveo recorded at station 1 (A) and 4 (B) in the Bari Canyon.

A

B

Fig. 5. Length-frequency distribution of

P. bogaraveo observed at station 1 (A) and 4

(B) in the Bari Canyon.



during offshore bottom longline fishing at depths shal-

lower than 400 m in the Southern Adriatic, of which the

hake (Merluccius merluccius) is the primary target species,

representing 40–70% of the total catch. The secondary

target species fished with noteworthy yields are the Euro-

pean conger (Conger conger) (10–20%), rockfish (H.

dactylopterus) (5–10%), blackspot seabream (P. bogarave-

o) (2–4%) while the shark species and wreckfish (P.

americanus) are rarely caught (De Zio et al. 1998; Ungaro

et al. 2005).

From experimental trawl catches carried out on the

slope (500–800 m) in the Southern Adriatic, Ungaro

et al. (1998a) reported the following abundance indices:

0.1 N�h�1 for C. conger; 2.5 N�h�1 for H. dactylopterus;

0.9 N�h�1 for M. merluccius. The abundance of these

three teleost fishes decreased markedly at depths >400 m.

Lepidopus caudatus and P. bogaraveo were only captured

on the shelf (Ungaro et al. 1998a).

The abundance of most species collected in this study

was also evaluated during the trawl surveys carried out as

part of the MEDITS project (Bertrand et al. 2002). In the

Southern Adriatic Sea, for the period 1994–99 and at a

depth range of 200–800 m, Spedicato et al. (2002)

reported density indices between 2 and 105 N�km�2 for

P. bogaraveo. For the period 1994–2009, Relini et al.

(2010) reported density indices between 0.6 and

0.8 N�km�2 for C. granulosus, 0.8 and 2.9 N�km�2 for

Dalatias licha, 39.5 and 274.3 N�km�2 for Etmopterus

spinax and 39.8 and 371.3 N�km�2 for Galeus melastomus.

Although the blackspot seabream is not among the

most abundant fish species in the Southern Adriatic Sea,

it was the most often observed fish with the highest num-

bers in the Bari Canyon. It is an appreciated fishery

resource mostly caught using longline and gillnet on the

slope (Morato et al. 2001; Chilari et al. 2006; Mytilineou

et al. 2013). Ontogenetic changes of habitat are character-

istic of this fish, with the juveniles appearing mainly in

shallower waters on muddy bottoms and larger fishes

often associated with offshore outcrops, banks and habi-

tats less accessible to trawling (Morato et al. 2001; Mene-

zes et al. 2006; Afonso et al. 2012; Mytilineou et al.

2013). Studies carried out using towed cameras and long-

line in the Santa Maria di Leuca (SML) cold-water coral

province showed this fish to be significantly associated

with the coral habitat (D’Onghia et al. 2011, 2012). The

residency of sub-adult and adult blackspot seabream at

the Condor seamount (Azores, mid-North Atlantic) was

ascertained using acoustic telemetry (Afonso et al. 2012).

Thus, the high numbers observed in the Bari Canyon

could be due to a preferential distribution of this fish in

A

B

Fig. 6. Fluctuations of visual count

(expressed as mean Max N with standard

deviation) and mean current velocity (ms�1

with standard deviation) at 443 m (A) and

577 m (B) in the Bari Canyon. Black bar:

night duration; horizontal line: MESOR of

visual count (9.8 in A and 3.2 in B).



a complex and heterogeneous habitat. Although the

lander was deployed on a sedimentary bioturbated sea

bed, P. bogaraveo is a very mobile fish species (Afonso

et al. 2012; Mytilineou et al. 2013) and at the depths and

areas where it was detected in the Bari Canyon, the exis-

tence of a megabenthic community dominated by a vari-

ety of cnidarians as major habitat-forming taxa, often in

association with sponges and serpulids has been revealed

using ROVs (Freiwald et al. 2009; Bo et al. 2012; Sanfi-

lippo et al. 2013; Angeletti et al. 2014).

The gregarious behavioural pattern known for blackspot

seabream is derived almost exclusively from fishery data

(Relini et al. 1999; Fabi et al. 2002, 2004; Chilari et al.

2006). Although specific patterns of shoaling and/or

schooling behaviour in the blackspot seabream have not

been investigated in detail here, at the depths where this

fish was observed, groups of individuals aggregate near the

bait, spending extensive amounts of time exploring the site

and remaining together. Schooling is considered an effi-

cient way of swimming which benefits the individual par-

ticipants in defeating and confounding predators, in faster

food localisation and more effective feeding as well as in

hydrodynamic advantages and in increasing reproductive

success (Pitcher 1993; Fr�eon & Misund 1999). The rhythms

in feeding activity in fish have often been explained under

various environmental conditions such as light level and

current speed (e.g. Lokkeborg et al. 2000 and references

therein; Doya et al. 2014 and references therein). At the

station where day–night variations in the visual counts

were investigated in the Bari Canyon, that is, at 577 m,

most fish were detected during the crepuscular period. This

is in agreement with Afonso et al. (2012) who detected

most fish during crepuscular and nocturnal periods over

the Condor seamount summits and reported that local

fishermen prefer to fish during these periods, avoiding day-

time fishing when fish are at deeper levels and less accessi-

ble. Further investigations are required to understand the

spatio-temporal variations in activity patterns of P. bogara-

veo in the Bari Canyon, as an increase in the visual counts

was also detected during afternoon at 443 m and when

water speed decreases at both stations. However, groups of

individuals of P. bogaraveo were observed to be attracted to

the bait, investigating, attacking and ingesting it both dur-

ing day and night hours. In the sharpsnout seabream (Dipl-

odus puntazzo), a fish species of the same Sparidae family,

both diurnal and nocturnal locomotor activities were

detected but feeding behaviour remained strictly diurnal

(Vera et al. 2006).

Although P. bogaraveo feeds both near the bottom on

benthic prey and in the water column on pelagic species

(Morato et al. 2001; Stergiou & Karpouzi 2002; Mytili-

neou et al. 2013), it can behave as a predator or scaven-

ger according its opportunistic encounters in theTable

3.Po

sition,reactionan

dactivity

byspeciesobserved

byMEM

Olander

intheBariCan

yon.

Species

Position

Reaction

Activity

Water

column

Near

seab

ed

On

seab

ed

No

reaction

Avoidan

ce-disturbed

Attraction

Actively

swim

ming

Quietly

swim

ming

Hovering

Moving

onseab

ed

Resting

onseab

edFeed

ing

Feed

ingbait

Chondrich

thyes

Cen

trophorusgranulosus

XX

X

Dalatiaslicha

XX

XX

Etmopterusspinax

XX

XX

Galeu

smelastomus

XX

XX

XX

X

Hexan

chusgriseus

XX

XX

Osteichthyes

Conger

conger

XX

XX

XX

X

Helicolenusdactylopterus

XX

XX

XX

XX

X

Hoplostethusmed

iterraneu

sX

XX

Lepidopuscaudatus

XX

XX

Merlucciusmerluccius

XX

XX

XX

Pagellusbogaraveo

XX

XX

XX

XX

X

Polyprionam

erican

us

XX

X



environment. The present study is the first in situ direct

observation of the variable feeding behaviour of P. bog-

araveo as well as of its gregarious habits.

Scavenger activity has also been observed in C. conger

which is considered a large opportunistic predator living

and foraging close to rocky areas, where it finds refuge

A B

Fig. 8. Individuals of P. bogaraveo actively swimming near the lander in a circle (A) and in a coordinated fashion (B).

Fig. 9. Individuals of P. bogaraveo feeding on bait.

A B C

Fig. 7. Individuals of Etmopetrus spinax (A), Conger conger (B) and Pagellus bogaraveo (C) feeding on bait.



during the day (Morato et al. 1999; Xavier et al. 2010).

Conger conger was collected in the Santa Maria di Leuca

coral area using fishing gears (D’Onghia et al. 2010) and

towed cameras, these latter revealing a swimming behav-

iour near the sea bed (D’Onghia et al. 2011). Using the

MEMO lander, the European conger was observed roam-

ing on the bottom and scavenging on the bait in the

SML cwc province (unpublished data). It could be

another species which would have a preferential distribu-

tion in structurally complex habitats like those built by

deep-sea corals. In this respect, Sulak et al. 2007 reported

Conger oceanicus burrowing into the base of Lophelia

bushes.

The teleost H. dactylopterus exhibited a clear behaviour-

al pattern of resting on the sea bed. This fish uses a wide

range of habitats, among which are cwc habitats and sub-

marine canyons, and is closely associated with the bottom

(e.g. Bourcier & Zibrowius 1973; Uiblein et al. 2003; Cos-

tello et al. 2005; Ross & Quattrini 2007; D’Onghia et al.

2010, 2011). Helicolenus dactylopterus seems to be a typical

sit-and-wait ambush predator feeding mainly on benthic

crustaceans and fish as well as on plankton organisms (Mi-

anzan et al. 1996; Nouar & Maurin 2000; Uiblein et al.

2003; Consoli et al. 2010). As it can also be a potential

prey of larger deep-sea predators, the observed behaviour

associated to its orange-red colour, undetectable to preda-

tors, can be considered a trade off between feeding strategy

and predator avoidance. Although this fish is frequently

caught using longlines, the observed behaviour and the

fact that it was never attracted by the bait do not make it a

typical scavenger fish.

The lower presence observed in the other common

species, such as M. merluccius, also deserves consideration

related to their distribution and behaviour. Even though

M. merluccius is among the most abundant demersal spe-

cies in the Southern Adriatic Sea (e.g. Marano et al. 1998;

Carlucci et al. 2009) the investigated sites in this study

are deeper than those where the species is mainly distrib-

uted. In fact, the greatest abundance of this fish is gener-

ally recorded on the shelf and shelf edge (e.g. Marano

et al. 1998; Carlucci et al. 2009). Most probably, M. mer-

luccius was only recorded during daytime because it

migrates up the water column during the night hours in

pursuit of pelagic prey (Carpentieri et al. 2005a,b),

Lepidopus caudatus individuals were also observed dur-

ing daylight hours feeding in the sediment, confirming

observations made in the Santa Maria di Leuca cwc prov-

ince (D’Onghia et al. 2011).

Adult individuals of P. americanus are usually solitary

swimmers and seem to have a preferential distribution

on larger lithoherms and hardgrounds (Messing et al.

1990) as well as in caves and on shipwrecks (Bianchi

et al. 1999). Using a longline and a baited lander,

respectively, the wreckfish was collected and observed in

the complex habitat of the Santa Maria di Leuca cwc

province (D’Onghia et al. 2012; Maiorano et al. in

press).

The deep-water velvet shark E. spinax is an active pred-

ator feeding on a wide range of benthic and free swim-

ming organisms, mostly represented by cephalopods

(Wurtz & Vacchi 1978; Carrass�on et al. 1992). E. spinax

was observed to investigate and feed on the bait in agree-

ment with observations by Gilat & Gelman (1984). These

authors reported that the diurnal distribution of E. spinax

was similar during the day and at night. During observa-

tions carried out in the SML cwc province using the same

baited lander, the bait was not consumed by the attracted

individuals of E. spinax (Sion et al. in press).

The shark G. melastomus has a diet based on epiben-

thic (mostly Calocaris macandreae) and benthopelagic

species (euphasids, ommastrephidae, Pasiphaea multiden-

tata, mesopelagic fishes) (Carrass�on et al. 1992). In the

Bay of Biscay G. melastomus was observed displaying a

high rate of forward locomotion close to the bottom but

no scavenger behaviour was observed (Uiblein et al.

2003).

With regard to C. granulosus, D. licha and H. griseus,

the present observations together with those carried out

in the Eastern and Central Mediterranean (Gilat & Gel-

man 1984; Jones et al. 2003; Sion et al. in press), indi-

cate that the dominant behaviour of deep-sea sharks

seems to be investigation at the foot of the lander with-

out ingesting the bait, suggesting that it is unlikely that

scavenging activity is an important feeding pattern for

these deep-sea sharks. Gilat & Gelman (1984) reported

that at 510 m C. granulosus was observed from noon to

late evening and was probably unaffected by fluctuations

in light. The small numbers of sharks observed in the

Bari Canyon could be related to their rare populations,

set at the top of the marine food web. However, the spo-

radic catches carried out in the Southern Adriatic Sea

have been suggested to be due to the intense fishing

pressure exercised on these k-selected deep-sea species

(Ungaro et al. 1996).

The presence of rare top predators, such as C. granulo-

sus and H. griseus as well as M. merluccius, which are

considered ‘vulnerable’ on the published IUCN Mediter-

ranean Regional Red List (Abdul Malak et al. 2011), and

large individuals of commercial species, such as P.

bogaraveo and M. merluccius, would suggest that this can-

yon, as observed in other marine regions (e.g. Yoklavich

et al. 2000; Farrugio 2012; Tecchio et al. 2013), acts as a

refuge area for several species impacted by fishing. How-

ever, all the above aspects require further investigations

to obtain additional information on fish distribution,

abundance and behaviour.



Acknowledgements

This study was carried out as part of the Italian RIT-

MARE project funded by the Italian Ministry of Univer-

sity and Research – MIUR. The authors acknowledge the

EU_7FP CoCoNet project. Thanks to Federica Foglini

and Franco Curci for the production of Fig. 1. Thanks

also to Daniela Potenza for the production of the figures

from video frames.

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Exploring composition and behaviour of fish fauna by in situ observations in the Bari Canyon...

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