Anali za istrske in mediteranske študije
Annali di Studi istriani e mediterranei
Annals for Istrian and Mediterranean Studies
Series Historia Naturalis, 33, 2023, 1
UDK 5                       Annales, Ser. hist. nat., 33, 2023, 1, pp. 1-142, Koper 2023  ISSN 1408-533X
KOPER 2023
Anali za istrske in mediteranske študije
Annali di Studi istriani e mediterranei
Annals for Istrian and Mediterranean Studies
Series Historia Naturalis, 33, 2023, 1
UDK 5                    ISSN 1408-533X 
                e-ISSN 2591-1783
ANNALES · Ser. hist. nat. · 33 · 2023 · 1
Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies
ISSN 1408-533X     UDK 5  Letnik 33, leto 2023, številka 1
e-ISSN 2591-1783
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ANNALES · Ser. hist. nat. · 33 · 2023 · 1
Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies
UDK 5                                                      Letnik 33, Koper 2023, številka 1                                      ISSN 1408-53 3X
e-ISSN 2591-1783 
VSEBINA / INDICE GENERALE / CONTENTS 2023(1)
BIOTSKA GLOBALIZACIJA
GLOBALIZZAZIONE BIOTICA
BIOTIC GLOBALIZATION
Andrea LOMBARDO
A New Mediterranean Record of the 
Sacoglossan Thuridilla mazda (Mollusca, 
Gastropoda) with a Review of its 
Distribution, Biology and Ecology ..........................
Nov sredozemski zapis o pojavljanju 
polža zaškrgarja vrste Thuridilla mazda 
(Mollusca, Gastropoda) s pregledom 
njene razširjenosti, biologije in ekologije
Deniz ERGUDEN, Sibel ALAGOZ ERGUDEN 
& Deniz AYAS On the Occurrence of Lutjanus 
argentimaculatus (Forsskål, 1775) in the 
South-Eastern Mediterranean, Turkey ..................
O pojavljanju mangrovskega rdečega hlastača 
Lutjanus argentimaculatus (Forsskål, 1775) v 
jugovzhodnem Sredozemskem morju (Turčija)
Adib SAAD, Lana KHREMA, Amina 
ALNESSER, Issa BARAKAT & 
Christian CAPAPÉ The First Substantiated 
Record of Areolate Grouper Epinephelus 
areolatus (Serranidae) and Additional 
Records of Pilotfish Naucrates ductor 
(Carangidae) from the Syrian Coast 
(Eastern Mediterranean Sea) ................................
Prvi potrjen zapis o pojavljanju rdečepikaste 
kirnje, Epinephelus areolatus (Serranidae), in 
dodatni zapis o pojavljanju pilota, Naucrates 
ductor (Carangidae), iz sirske obale (vzhodno 
Sredozemsko morje)
Okan AKYOL & Vahdet UNAL
Additional Record of Sillago suezensis 
(Sillaginidae) from the Aegean Sea, Turkey ..........
Nov zapis o pojavljanju rdečemorskega 
mola Sillago suezensis (Sillaginidae) 
v turškem Egejskem morju
SREDOZEMSKI MORSKI PSI 
SQUALI MEDITERRANEI
MEDITERRANEAN SHARKS
Hakan KABASAKAL, Uğur UZER & 
F. Saadet KARAKULAK 
Occurrence of Deep-Sea Squaliform 
Sharks, Echinorhinus brucus 
(Echinorhinidae) and Centrophorus 
uyato (Centrophoridae), 
in Marmara Shelf Waters .........................................
Pojavljanje dveh globokomorskih 
morskih psov Echinorhinus brucus 
(Echinorhinidae) in Centrophorus 
uyato (Centrophoridae), 
v vodah Marmarskega šelfa
Khadija OUNIFI-BEN AMOR, 
Mohamed Mourad BEN AMOR, 
Marouène BDIOUI & 
Christian CAPAPÉ 
Additional Captures of Smoothback 
Angel Shark Squatina oculata (
Squatinidae) from the Tunisian Coast ...................
(Central Mediterranean Sea)
Nova ulova pegastega sklata 
Squatina oculata (Squatinidae) 
iz tunizijske obale (osrednje 
Sredozemsko morje)
Alessandro DE MADDALENA, 
Marco Giovanni BONOMO, 
Andrea CALASCIBETTA & 
Lorenzo GORDIGIANI
On a Large Shortfin Mako 
Shark Isurus oxyrinchus (Lamnidae) 
Observed at Pantelleria 
(Central Mediterranean Sea) ..................................
O velikem primerku atlantskega 
maka, Isurus oxyrinchus (Lamnidae), 
opaženega blizu Pantellerie 
(osrednje Sredozemsko morje)
27
37
43
1
13
7
19
ANNALES · Ser. hist. nat. · 33 · 2023 · 1
IHTIOFAVNA
ITTIOFAUNA
ICHTHYOFAUNA
Christian CAPAPÉ, Christian REYNAUD & 
Farid HEMIDA The First Well-Documented 
Record of Maltese Skate Leucoraja 
melitensis (Rajidae) From the Algerian 
Coast (Southwestern Mediterranean Sea) ............
Prvi potrjeni primer o pojavljanju skata vrste 
Leucoraja melitensis (Rajidae) iz alžirske obale 
(jugozahodno Sredozemsko morje)
Alessandro NOTA, Sara IGNOTO, Sandro 
BERTOLINO & Francesco TIRALONGO
First Record of Caranx crysos (Mitchill, 1815) in 
the Ligurian Sea (Northwestern Mediterranean Sea) 
Suggests Northward Expansion of the Species ...........
Prvi zapis o pojavljanju modrega trnoboka 
Caranx crysos (Mitchill, 1815) v Ligurskem 
morju (severozahodno Sredozemsko morje) 
dokazuje širjenje vrste proti severu
Alen SOLDO The First Marine Record of 
Northern Pike Esox lucius Linnaeus, 1758 
in the Mediterranean Sea .......................................
Prvi morski zapis o pojavljanju ščuke Esox lucius 
Linnaeus, 1758 v Sredozemskem morju
Mourad CHÉRIF, Rimel BENMESSAOUD, 
Sihem RAFRAFI-NOUIRA & Christian CAPAPÉ
Diet and Feeding Habits of the Greater Weever 
Trachinus draco (Trachinidae) from the Gulf of 
Tunis (Central Mediterranean Sea) ........................
Prehranjevalne navade morskega zmaja 
Trachinus draco (Trachinidae) iz Tuniškega 
zaliva (osrednje Sredozemsko morje)
Laith A. JAWAD & Okan AKYOL
Skeletal Abnormalities in a Sphyraena 
sphyraena (Linnaeus, 1758) and a Trachinus 
radiatus Cuvier, 1829 Collected from the 
North-Eastern Aegean Sea, Izmir, Turkey ...........
Skeletne anomalije na primerkih vrst Sphyraena 
sphyraena (Linnaeus, 1758) in Trachinus 
radiatus Cuvier, 1829, ujetih v severovzhodnem 
Egejskem morju (Izmir, Turčija)
Deniz ERGUDEN, Sibel ALAGOZ ERGUDEN & 
Deniz AYAS A Rare Occurrence and Confirmed 
Record of Scalloped Ribbonfish Zu cristatus 
(Osteichthyes: Trachipteridae) in the Gulf 
of Antalya (Eastern Mediterranean), Turkey ..........
O redkem pojavljanju in potrjeni najdbi 
čopaste kosice Zu criistatus (Osteichthyes: 
Trachipteridae) v Antalijskem zalivu (vzhodno 
Sredozemsko morje), Turčija
FAVNA
FAUNA
FAUNA
Nicola BETTOSO, Lisa FARESI, Ida Floriana 
ALEFFI & Valentina PITACCO
Epibenthic Macrofauna on an Artificial Reef 
of the Northern Adriatic Sea: a Five-Years 
Photographic Monitoring ......................................
Epibentoška makrofavna na umetnem 
podvodnem grebenu v severnem Jadranu: pet 
letni fotografski monitoring
Roland R. MELZER, Martin PFANNKUCHEN, 
Sandro DUJMOVIČ, Borut MAVRIČ & 
Martin HEß First Record of the Golden 
Coral Shrimp, Stenopus spinosus Risso, 
1827, in the Gulf of Venice ...................................
Prvi zapis o pojavljanju koralne kozice, 
Stenopus spinosus Risso, 1827, 
v Beneškem zalivu
Abdelkarim DERBALI, Nour BEN MOHAMED & 
Ines HAOUAS-GHARSALLAH
Age, Growth and Mortality of 
Surf Clam Mactra stultorum 
in the Gulf of Gabes, Tunisia .................................
Starost, rast in smrtnost koritnice Mactra 
stultorum v Gabeškem zalivu (Tunizija)
Cemal TURAN, Servet Ahmet DOĞDU & 
İrfan UYSAL
Mapping Stranded Whales in 
Turkish Marine Waters ..........................................
Popisovanje nasedlih kitov v 
turških morskih vodah
OBLETNICE
ANNIVERSARI
ANNIVERSARIES
Martina ORLANDO-BONACA & Patricija 
MOZETIČ
Šestdeset let morskega biologa Lovrenca Lipeja .....
Kazalo k slikam na ovitku ...................................
Index to images on the cover ..............................
113
119
141
51
75
55
61
67
89
99
127
139
141
ANNALES · Ser. hist. nat. · 33 · 2023 · 1
ANNALES · Ser. hist. nat. · 33 · 2023 · 1
99
received: 2023-04-03                 DOI 10.19233/ASHN.2023.14
EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN 
ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC MONITORING
Nicola BETTOSO, Lisa FARESI & Ida Floriana ALEFFI
Agenzia Regionale per la Protezione dell’Ambiente del Friuli Venezia Giulia (ARPA FVG), via Cairoli 14 – 33053 Palmanova (UD), Italy
e-mail: nicola.bettoso@arpa.fvg.it
Valentina PITACCO
Marine Biology Station Piran, National Institute of Biology, Fornače 41, 6330 Piran, Slovenia
ABSTRACT
Artificial reefs (ARs) are man-made structures used with the aim of improving fisheries and increasing natural 
production of biological resources. In 2006 an AR made of three types of modules was sunk near an underwater 
sewage outfall. The objectives of the project were: (a) to use the AR to restock some target species of commercial 
interest and (b) to promote biodiversity in selected areas. The epibenthic macrofauna that had settled on this AR 
was annually monitored for five years (2007- 2011) using non-destructive photographic methods. A total of 88 
taxa from 8 phyla were identified, with a predominance of Porifera, Mollusca and Ascidiacea. Among the types of 
modules used to construct the AR, polyethylene panel nets were functional for bivalve settlement in the first year, 
whereas concrete structures seemed to perform best in promoting biodiversity in terms of species richness in the 
long term. Nevertheless, the 5-year monitoring period was too short to speculate on the stability or homeostasis of 
communities settled on the AR in terms of ecological succession.
Key words: artificial reefs, macrozoobenthos, Adriatic Sea, photographic monitoring
LA MACROFAUNA EPIBENTONICA DI UNA BARRIERA ARTIFICIALE SOMMERSA 
DELL’ALTO ADRIATICO: RISULTATI DI UN MONITORAGGIO FOTOGRAFICO 
QUINQUENNALE
SINTESI
Le barriere artificiali (BA) sono strutture solitamente utilizzate per incrementare le rese di pesca. Nel 2006 è 
stata realizzata una BA, costruita con tre diverse tipologie di moduli e situata in prossimità di un dotto fognario 
sottomarino. Gli obiettivi del progetto erano: (a) sperimentare la BA per il ripopolamento di alcune specie di 
interesse commerciale e (b) promuovere la biodiversità. La macrofauna epibentonica insediata su questa BA è 
stata monitorata a cadenza annuale durante 5 anni (2007-2011) per mezzo di rilievi fotografici. Complessiva-
mente sono stati identificati 88 taxa appartenenti ad 8 phyla, in cui i Porifera, Mollusca ed Ascidiacea sono stati 
prevalenti. I pannelli in rete di polietilene sono stati efficaci durante il primo anno per l’insediamento dei molluschi 
bivalvi, mentre nel lungo periodo il cemento si è dimostrato il più efficace per promuovere la biodiversità in 
termini di ricchezza specifica. Il monitoraggio di 5 anni comunque non è stato del tutto soddisfacente per trarre 
conclusioni sulla successione ecologica della comunità insediata sulla BA.
Parole chiave: Barriere artificiali, macrozoobenthos, mare Adriatico, monitoraggio fotografico
ANNALES · Ser. hist. nat. · 33 · 2023 · 1
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Nicola BETTOSO et al.: EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC ..., 99–112
INTRODUCTION
Artificial reefs (ARs) are man-made structures 
that have long been used with the aim of improving 
fisheries by concentrating fish and increasing the 
natural production of biological resources (Bohn-
sack & Sutherland, 1985). Parenzan (1957) was one 
of the first scientists to suggest the use of artificial 
structures as a tool to increase fishery production 
in oligotrophic environments by sinking wrecks and 
testing ad hoc experimental areas (Parenzan, 1986). 
In Italy the first AR was made in December 1970, 
when a team of recreational fishermen, without any 
scientific support, obtained the permission to sink 
1,300 cars at depth between 35 and 50 m in order 
to discourage trawling and to improve recreational 
fishing (Relini & Orsi Relini, 1989). The first scientif-
ically oriented AR was built in 1974 in the Adriatic 
Sea, with stones and concrete blocks (Bombace, 
1989). After about 40 years, Fabi et al. (2011) report-
ed more than 70 ARs along the Italian coast, most 
of them built thanks to the financial support of the 
European Community. 
In the oligotrophic waters of the Mediterranean, 
ARs have been used mainly to protect Posidonia 
oceanica meadows from illegal trawling and to 
increase habitat complexity and species diversity 
(Relini et al., 1994; Gonzalez-Correa et al., 2005; 
Ponti et al., 2015). In contrast, in the eutrophic wa-
ters of the central and northern Adriatic, ARs have 
been used to increase fishery yields (Bombace et 
al., 1994; Bombace et al., 1997). Concrete, pebbles, 
limestone rocks, and PVC are the most common 
materials used to build ARs (Toledo et al., 2020 
and references therein). The concrete is frequently 
used because it is cheap and allows the realization 
of different structures in terms of shape and size. 
In addition, concrete is resistant to chemical and 
physical marine actions, thus ensuring long duration 
(Fabi et al., 2011; Ponti et al., 2015).
ARs are intentionally placed on the seafloor with 
the goal of imitating the function of a natural reef, 
therefore most studies after the installation of an AR 
are focused on the ecological succession of benthic 
organisms on these manmade substrates (Toledo et 
al., 2020 and references therein). The colonization 
process can be broadly divided in early and late 
succession: the former is associated with the first or-
ganisms that settle on the ARs, including organized 
microorganisms creating microbial mats or biofilms; 
the latter is characterized by the arrival of more 
complex organisms, which in turn attract predator 
species from further up the trophic chain (Herbert et 
al., 2017; Toledo et al., 2020 and references therein).
The AR analyzed in the present study was based 
on guidelines developed as part of the project ADRI.
BLU (2006). This project addressed the sustainable 
management of fisheries activities and resources in 
the northern Adriatic Sea. More information on the 
project is available on internet in Italian. The specif-
ic objectives of the project were: (a) to use the AR in 
Fig. 1: Map of the study area, the circle indicates the 
terminal tract of Lignano Sabbiadoro sewage outfall. 
Sl. 1: Zemljevid obravnavanega območja z označenim 
delom terminala kanalizacije mesta Lignano Sabbia-
doro (krogec). 
Fig. 2: Particular of the terminal tract of the sewage out-
fall, location of the AR units and photo sampling points.
Sl. 2: Del terminala kanalizacije, lega UPG in 
vzorčevalne postaje fotografskega monitoringa. 
101
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Nicola BETTOSO et al.: EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC ..., 99–112
order to restock some target species of commercial 
interest, such as pectinid bivalves, and (b) to pro-
mote biodiversity in some selected areas, such the 
Gulf of Trieste for the Region Friuli-Venezia Giulia. 
In this context, in 2006, an AR was sunk three miles 
off Lignano Sabbiadoro, near an underwater sewage 
outfall (Solis-Weiss et al., 2007, and references 
therein), and the settled macrofauna was monitored 
annually for the following five years (2007- 2011) 
using photographic methods based on a guide spe-
cifically developed within this project (Arpa FVG, 
2007; Bettoso et al., 2023). The photographic meth-
od is less accurate than other sampling techniques, 
but has the advantage of being non-destructive, 
fast, and efficient enough to monitor the evolution 
of epibenthic communities on the AR with a good 
benefit-cost ratio. In the present work the results of 
this photographic monitoring are presented.
MATERIAL AND METHODS
Study area
The AR was located in the Gulf of Trieste, the 
northernmost part of the Adriatic Sea (Fig. 1). It is 
a shallow semi-enclosed basin (max depth 25 m), 
characterized by the largest tidal amplitudes and 
the lowest winter temperatures in the Mediterra-
nean Sea (Boicourt et al., 1999), high temperature 
and salinity variations, and important stratification 
of the water column (Stravisi, 1983). The hydrody-
namism is related mainly to the ascending eastern 
current coming from the Istrian coast. The general 
circulation pattern is predominantly counterclock-
wise in the lower layer and clockwise in the surface 
layer. This circulation, especially in the surface 
layer can be modulated by prevailing winds from 
Fig. 3: a) Tecnoreef ® pyramid (TR); b) Ecomare pyramids (EM); c) Polyethylene net for scallops settlement (PE) 
(photos: Arpa FVG).
Sl. 3: a) Tecnoreef ® piramida (TR); b) Ecomare piramide (EM); c) Polietilenska mreža za naseljevanje pokrovač 
(PE) (fotografije: Arpa FVG).
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Nicola BETTOSO et al.: EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC ..., 99–112
eastern quadrants such as Bora (Stravisi, 1983). 
The sediments are quite varied, from sands with 
beachrocks to muds, predominantly detritical, 
and the associated biocoenoses of the Gulf belong 
mainly to the DC (Détritique Côtier), DE (Détri-
tique Envasé) and VTC (Vases Terrigènes Côtières) 
biocoenoses, as defined by Pérès & Picard (1964) 
(Solis-Weiss et al., 2001). 
The AR was deployed in August/September 2006 
about 3 nautical miles off Lignano Sabbiadoro near the 
underwater sewage outfall (Fig. 1), where anchoring and 
fishery are forbidden. The site is located at 16 m depth, 
where the sediment consists of pelitic sand and very 
sandy pelite, and benthic community of the soft bottom 
is mainly represented by the biocoenosis of Coastal De-
tritic (DC) (Solis-Weiss et al., 2007). The AR consisted of 
Fig. 4: Taxa richness (TR, Tecnoreef; EM, ecomare; PE, polyethylene). 
Sl. 4: Pestrost taksonov (TR, Tecnoreef; EM, ecomare; PE, polietilen-
ska mreža). 
Fig. 5: a) K-dominance curves for Tecnoreef (TR), b) K-dominance curves for ecomare (EM); c) K-dominance curves 
for polyethylene net (PE).
Sl. 5: Krivulje K-dominance za Tecnoreef (TR), b) krivulje K-dominance za ecomare (EM); c) krivulje K-dominance za 
polietilensko mrežo (PE).
103
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Nicola BETTOSO et al.: EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC ..., 99–112
Tab. 1: Taxa richness year by year on every AR module (TR – Tecnoreef, EM – ecomare, PE – polyethylene net) 
and Mann-Kendall test (ns – not significant, in bold – upward trend).
Tab. 1: Pestrost taksonov v posameznih letih na vsakem UPG modulu (TR – Tecnoreef, EM – ecomare, PE – 
polietilenska mreža) in Mann-Kendallov test (ns – ni statistično značilen, mastni tisk – trend naraščanja).
TR 2007 2008 2009 2010 2011 Mann-Kendall
tot 19 31 49 41 46 ns
Porifera 1 6 13 11 16 0.042
Mollusca 8 8 10 7 9 ns
Echinodermata 5 2 3 5 2 ns
Cnidaria 0 2 3 2 3 ns
Chordata 3 9 12 12 9 ns
Bryozoa 0 1 2 2 3 0.0083
Arthropoda 0 1 3 0 1 ns
Annelida 2 2 3 2 3 ns
EM 2007 2008 2009 2010 2011
tot 12 23 32 33 37 0.0083
Porifera 2 4 8 7 8 0.042
Mollusca 7 5 8 7 10 ns
Echinodermata 0 2 3 3 2 ns
Cnidaria 0 1 2 2 3 0.0083
Chordata 2 8 7 9 5 ns
Bryozoa 0 1 2 2 3 0.0083
Arthropoda 0 1 0 1 2 ns
Annelida 1 1 2 2 4 0.042
PE 2007 2008 2009 2010 2011
tot 27 17 25 30 30 ns
Porifera 1 1 7 7 11 0.042
Mollusca 12 3 6 8 5 ns
Echinodermata 3 0 1 2 2 ns
Cnidaria 2 3 3 2 3 ns
Chordata 6 6 5 6 4 ns
Bryozoa 0 1 1 2 2 0.042
Arthropoda 1 0 0 1 0 ns
Annelida 2 3 2 2 3 ns
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Nicola BETTOSO et al.: EPIBENTHIC MACROFAUNA ON AN ARTIFICIAL REEF OF THE NORTHERN ADRIATIC SEA: A FIVE-YEARS PHOTOGRAPHIC ..., 99–112
two different units (hereafter AR units) placed on both 
sides of the sewage duct, at a distance of 500 m from 
the end section of the sewer (Fig. 2). These AR units 
were made of three different types of artificial modules 
(hereafter AR modules) placed in a symmentrical and 
specular way with respect to the sewer (Fig. 2). These 
modules were: 30 pyramids made of concrete slabs (TR 
– declared “sea-friendly” by the producer Tecnoreef®, 
manifactured using only natural components, without 
synthetic additives; pH – 9; 180 cm height) (Fig. 3a); 4 
pyramids made of PVC tubes (EM – Ecomare, diameter 
of tubes is 100 cm and pyramid high is 273 cm) (Fig. 
3b); 4 sets of polyethylene panel nets (PE) (mesh size 40 
mm, length 100 m, height 300 cm, width 100 cm) used 
for the settlement of scallop larvae (Fig. 3c). 
Sampling and analysis
Epibenthic faunal communities on the AR were 
surveyed once a year using SCUBA photo-sampling 
(Fujifilm Finepix F30 model). Pictures of a 50x50 cm 
(2,500 cm2) quadrat (divided in 4 subquadrats) were 
taken at 16 points for each unit (4 for TR, 4 for EM, and 
8 for PE) for a total of 32 sampling points (displayed 
in Fig. 2). Animals that were readily identifiable in the 
photographs were determined to the lowest possible 
taxonomic level, whereas those not easily recogniz-
able at the species level were collected and stored for 
determination at the laboratory (Bettoso et al., 2023). 
The first survey took place in June 2007, nine months 
after the sinking of the AR, and was repeated each 
spring/summer until 2011, for a total of 132 quadrats 
analysed. Cluster, K-dominance curves and SIMPER 
analysis (Clarke and Warwick, 2001) were performed 
on presence/absence data subdivided for type of AR 
module and year of sampling. Mann Kendall test was 
applied to statistically assess if there is a monotonic 
upward or downward trend of the variable of interest 
over time.
RESULTS
A total of 88 taxa of epibenthic invertebrates, be-
longing to 8 phyla, were detected by photo sampling, 
of which 75 were recorded on concrete pyramids 
(TR), 57 in both PVC pyramids (EM) and polyethylene 
panel nets (PE). The most abundant phyla in terms 
of taxa richness were Mollusca followed by Porifera 
and Chordata, the latter represented by class Asci-
diacea. The same pattern was observed considering 
the three types of AR modules separately, except for 
TR, where Porifera slightly outweigh Mollusca (Fig. 
4). The highest taxa richness was observed on TR 
modules for all phyla except for Cnidaria, showing 
the highest richness on PE modules (Fig. 4). Con-
sidering the total number of taxa, an increase with 
time was observed only for EM (Mann-Kendall test, 
Tab. 1). Considering the different phyla separately, 
Porifera and Bryozoa were the only groups to show 
an increasing trend in richness values in all types of 
AR modules (Tab. 1).
The K-dominance curves generated for TR 
showed decreasing dominance from 2007 to 2008, 
whereas curves for 2009, 2010 and 2011 showed 
a comparable shape (Fig. 5a). Similar pattern was 
observed for EM, although the curves from 2009 
to 2011 intersect at one point (Fig. 5b). For PE, the 
shape of the curve for 2007 was more similar to 
the curves for 2009-2011, whereas the curve for 
2008 was the steepest and showed the highest 
dominance (Fig. 5c).
The dendrogram clearly separated the first 
survey of 2007 (Group A: EM07, TR07, PE07) at a 
similarity level of about 20%, whereas the Group 
B included TR08 and EM08 surveyed in 2008. The 
remaining clusters (2009-2011) were grouped at a 
similarity level of about 60%, except PE08 which 
was grouped apart (Group C). The cluster 2009-
2011 was subdivided on the basis of the type of 
Fig. 6: Dendrogram on the basis of years (07-11) and AR module (TR, EM, PE).
Sl. 6: Dendrogram na podlagi različnih let (07-11) in modula AR (TR, EM, PE). 
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Tab. 2: SIMPER analysis performed on clusters of the dendrogram (cut off 50%).
Tab. 2: SIMPERjeva analiza posameznih gruč na dendrogramu (rez na nivoju 50%). 
  Group A 2007 (TR, EM, PE)   Group B 2008 (TR, EM)   Group C 2008 (PE)
POR Crambe crambe POR Crambe crambe CNI Cereus pedunculatus
MOL Hexaplex trunculus POR Ulosa digitata ANN Serpulidae indet.
MOL Ostrea edulis MOL Hexaplex trunculus CHO Phallusia mammillata
MOL Mimachlamys varia MOL Ostrea edulis CHO Styela plicata
MOL Pinna nobilis ANN Serpulidae indet. CHO Didemnum lahillei
MOL Mytilus galloprovincialis CHO Didemnum lahillei  
MOL Flexopecten glaber CHO Phallusia mammillata  
ANN Serpulidae indet. CHO Phallusia fumigata  
CHO Phallusia mammillata CHO Diplosoma listerianum  
CHO Botryllus schlosseri  
  Group D 2009-2011 (TR) Group E 2009-2011 (EM) Group F 2009-2011 (PE)
POR Crambe crambe POR Crambe crambe POR Crambe crambe
POR Dysidea fragilis POR Dysidea fragilis POR Ulosa digitata
POR Haliclona (Reniera) mediterranea POR
Haliclona (Reniera) 
mediterranea POR Dysidea fragilis
POR Phorbas fictitius POR Ulosa digitata POR Haliclona (Reniera) mediterranea
POR Dictyonella incisa CNI Cereus pedunculatus CNI Epizoanthus sp.
POR Ulosa digitata MOL Hexaplex trunculus CNI Cereus pedunculatus
CNI Cereus pedunculatus MOL Ostrea edulis MOL Hexaplex trunculus
MOL Hexaplex trunculus MOL Anomia ephippium MOL Ostrea edulis
MOL Ostrea edulis MOL Mimachlamys varia MOL Anomia ephippium
MOL Mimachlamys varia ANN Serpulidae indet. MOL Mytilus galloprovincialis
ANN Serpulidae indet. BRY Bryozoa indet. ANN Serpulidae indet.
BRY Bryozoa indet. BRY Schizobrachiella sanguinea BRY Bryozoa indet.
BRY Schizobrachiella sanguinea ECH Ocnus planci ECH Ophiothrix fragilis
ECH Ocnus planci CHO Phallusia mammillata CHO Didemnum lahillei
CHO Microcosmus sp. CHO Phallusia fumigata CHO Diplosoma listerianum
CHO Didemnum lahillei CHO Didemnum lahillei  
CHO Diplosoma listerianum CHO Microcosmus sp.    
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AR modules rather than years (Group D: TR09, 
TR10, TR11; Group E: EM09, EM10, EM11; Group 
F: PE09, PE10, PE11) (Fig. 6). 
SIMPER analysis was used to identify the most 
representative species for each cluster of the dendro-
gram (Tab. 2). Crambe crambe was the first Porifera 
species recorded on the AR and was found every year 
and on all types of AR modules, except PE in 2008. 
In the second year, Ulosa digitata was detected on TR 
and EM and was observed on all types of AR modules 
in all subsequent years. Dysidea fragilis and Haliclo-
na (Reniera) mediterranea were among characteristic 
species from 2009 onwards. SIMPER identified two 
more species of Porifera only on TR (group D) (Tab. 
2). Considering phylum Annelida, the taxon Serpuli-
dae was constantly found in every year and all types 
of AR modules. The phylum Mollusca was mainly 
represented by bivalves and had the highest number 
of characteristic species in 2007. Flexopecten glaber 
and Pinna nobilis were recorded only during the first 
year, whereas the gastropod Hexaplex trunculus and 
the oyster Ostrea edulis characterized all groups of 
dendrogram, except the group C (PE in 2008) where 
no molluscs were identified by SIMPER analysis. 
Mytilus galloprovincialis was included in the group F 
(PE 2009-11) and Mimachlamys varia in those D and 
E (TR and EM 2009-11). The phylum Cnidaria was 
represented mainly by Cereus pedunculatus, being 
characteristic of group C on PE in 2008 and in all types 
of AR modules from 2009 to 2011. The Chordata As-
cidiacea were characteristic in all cluster groups. It is 
interesting to note the exclusive presence of Botryllus 
schlosseri in 2007 and the particular abundance of 
Phallusia mammillata in 2007 and 2008 on all types 
of AR. Didemnum lahillei was among characteristic 
species from 2008 onwards (Tab. 2).
DISCUSSION AND CONCLUSIONS
The Gulf of Trieste is characterized by various 
environmental and anthropogenic pressures that affect 
benthic communities, such as periodic “mare sporco” 
phenomena (mucilage aggregations), episodes of hy-
poxia and anoxia, significant riverine inflow, intense 
maritime traffic, intensive fishery, mariculture and 
others (Stachowitsch & Fuchs, 1995; Solis-Weiss et al., 
2004). These pressures could lead to changes in the 
soft- and hard-bottom benthic communities, at least in 
terms of succession (Mavrič et al., 2010). 
A recent census of ARs in the Adriatic Sea recorded 
a total of 47 sites along the Italian coast, 8 of which 
are located in the Gulf of Trieste (Minelli et al., 2021). 
The oldest AR of the gulf was built in 1978 inside the 
Miramare Marine Protected Area in Trieste, while the 
last one was established near mussel farms on the 
maritime border Italy-Slovenia (Project EcoSea, 2016). 
The site for the present work, near the sewage outfall 
off Lignano Sabbiadoro was selected to avoid damage 
to the AR and sewage pipeline from anchoring and 
fishing, and because this area is characterized by the 
presence of pelitic sand or very sandy pelite. In fact, 
this sediment texture is suitable to prevent the sinking 
Fig. 7: Tecnoreef pyramid (TR) in 2015 (photo: G. Pessa).
Sl. 7: Tecnoreef piramida (TR) leta 2015 (foto: G. Pessa).
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of AR modules into the seafloor and the accumulation 
of muddy sediment on the AR, which could inhibit 
the development and diversification of epibenthic 
organisms. In addition, the proximity of natural rocky 
outcrops (locally knows as Trezze) very rich in term 
of epibenthic fauna (Lipej et al., 2016; Bettoso et al., 
2023), could represent a good larval source for the set-
tlement of species common in coralligenous biotopes. 
According to Ponti et al. (2015) the environmental 
conditions are the main drivers of the recruitment 
processes on ARs. In particular, the sedimentation 
rate seemed the most important in the establishment 
of different benthic assemblages and therefore in the 
ecological effectiveness of the ARs (Ponti et al., 2015). 
In the present study the area off Lignano Sabbiadoro 
shows a heterogeneous sediment texture: AR modules 
are located on prevailing sandy sediment close to 
the rocky outcrops, but just 1 Nm toward the coast 
the sediment becomes muddy due to the pelitic belt 
originated by the Tagliamento river mouth. Thus, this 
zone can be considered a transitional habitat between 
areas with high sedimentation rate and coralligenous 
biotopes.
A total of 88 taxa belonging to 8 phyla, identified 
for the present work on the AR by photographic 
monitoring only, is a high number when compared 
to the 196 taxa of epibenthic invertebrates recorded 
by Bettoso et al. (2023) using the same method on 45 
natural rocky outcrops not far from the study area. 
Porifera, Mollusca and Ascidiacea predominated on 
the AR, as well as on those natural rocky outcrops 
(Bettoso et al., 2023). The faunal component on the 
AR dominated over macroflora, because the site was 
at 16 m depth with recurring events of water turbidity. 
The same was observed for the natural rocky outcrops 
(Bettoso et al., 2023). So, the proximity of the sewage 
pipeline did not seem to have a negative impact on 
the epibenthic fauna settled on the AR. Consistently, 
no negative impact had been observed on the soft 
bottom macrofauna near the same sewage outfall by 
Solis-Weiss et al. (2007).
According to Ponti et al. (2015) the shape and ma-
terials of ARs were of little importance in determining 
the structure of the benthic assemblage. Conversely, in 
the present work, some differences between the types 
of AR modules deployed were observed. In particular, 
TR showed overall higher richness values compared to 
EM and PE (Fig. 4). 
The cluster showed that those differences are not 
evident immediately after the deployment of the AR (in 
2007 all AR modules are clustered together), but from 
2009 on, when benthic communities are clustered 
by type of modules rather than by years. On TR and 
EM total taxa richness increased from 2007 to 2008, 
mainly due to Porifera and Ascidiacea. From the third 
year (2009) the number of taxa did not increase on any 
AR module (Tab. 1). Dominance decreased for 2007 to 
2009, and did not changed in the subsequent years on 
both TR and EM. Unfortunately the photographic mon-
itoring on epibenthic fauna lasted only 5 years and 
further observations on the variability and/or stability 
on this community was not possible. Nevertheless, 
some pictures taken in 2015 on TR modules showed 
an assemblage with oysters, sponges and tunicates 
clearly observable (Fig. 7). 
In the first year of monitoring the richness of bi-
valves was much higher on polyethylene panel nets 
(PE) than on concrete pyramids (TR) and PVC pyramids 
(EM). However, the permanent immersion of PE did not 
allow similar settlement of scallops and other bivalve 
Fig. 8: Smooth scallops (Flexopecten glaber) and 
Spirobranchus triqueter settled on polyethylene nets 
(PE) (photo: Arpa FVG).  
Sl. 8: Gladka pokrovača (Flexopecten glaber) in 
Spirobranchus triqueter sta se naselila na polietilensko 
mrežo (PE) (foto: Arpa FVG).  
Fig. 9: Smooth scallops (F. glaber) on sea floor nearby 
polyethylene nets (photo: Arpa FVG).  
Sl. 9: Gladka pokrovača (F. glaber) na morskem dnu v 
bližini polietilenskih mrež (foto: Arpa FVG).  
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in subsequent years as in 2007. On PE a decrease in 
richness was observed from 2007 to 2008 (Tab. 1), and 
at the same time dominance increased (Fig. 5c). 
Polyethylene panel nets (PE) were specifically 
deployed to test the settlement of scallops, particularly 
for the smooth scallop Flexopecten glaber. This species 
is a very appreciated fisheries resource in the northern 
Adriatic Sea. It is usually caught together with Pecten 
jacobaeus by rapido, a type of beam trawl, used only 
in Adriatic Sea, with a very severe impact on benthic 
communities and whose use should be better regulated 
(Giovanardi et al., 1998). The effectiveness of panel nets 
or collector bags as substratum for the settlement of F. 
glaber pediveligers has been successfully tested in other 
areas of the Gulf of Trieste (Orel & Zamboni, 2003). A 
massive production of this resource was found on PE 
also in the present study (Fig. 8). After 10 months from 
the immersion of 400 m of these panel nets, the smooth 
scallops reached a commercial size with an average 
density on the seafloor of 17 ind. m-2 and an estimated 
total biomass of 6,597 kg (Fig. 9) (Arpa FVG, 2007). 
Nevertheless, the effectiveness of panel nets for larval 
settlement of the smooth scallops, the queen scallop 
Aequipecten opercularis and other bivalves requires a 
clean substrate and a correct period of immersion for 
each species in order to detect the massive swarming of 
larvae (Orel & Zamboni, 2003). For instance, the period 
between June and September was considered the best 
for the captation of the variegated scallop Mimachlamys 
varia in the central Adriatic (Marguš et al., 1993).
PE proved to be efficient also for the settlement of 
the critically endangered species Pinna nobilis, but 
almost all individuals were observed in 2007. So even 
if the species settles successfully, it probably does not 
survive in such types of structures. ARs made of plastic 
nets are recommented as larval captators by IUCN 
(Kersting & Hendriks, 2019) and have been extensively 
studied and used in different areas of the Mediterra-
nean since the first mass mortality event of P. nobilis 
in 2016 (Kersting et al., 2020). According to IUCN 
guidelines the main reproduction period of P. nobilis is 
from May to August and the main settlement period is 
between July and September (in the western Mediter-
ranean), so collectors are usually deployed in June and 
Fig. 10: Macrofauna assemblage on concrete plates (TR) in 2009 with the presence of the sponge Haliclona 
(Reniera) mediterranea (photo: Arpa FVG).
Sl. 10: Združba makrofavne na betonskih ploščah (TR) v letu 2009 s spužvo Haliclona (Reniera) mediterranea 
(foto: Arpa FVG).
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removed in October-November (Kersting & Hendriks, 
2019). These periods could change depending on 
environmental conditions (e.g., water temperature) in 
different years and Mediterranean regions (Kersting & 
García-March, 2017). 
Based on the present work it is not possible to de-
fine a clear species succession, although, as the results 
show, some taxa such as Serpulidae, Ostrea edulis 
and Crambe crambe characterized the community 
every year; other like Botryllus schlosseri and F. glaber 
characterized only the first year, while some Porifera 
species (e. g. Dysidea fragilis and Haliclona (Reniera) 
mediterranea) characterized the community from 
2009 onward (Fig. 10; SIMPER analysis). Serpulids, 
like Spirobranchus triqueter, are considered among pi-
oneer species (Maggiore & Keppel, 2006; Nicoletti et 
al., 2007) able to drive the early stages of settlement, 
because of encroaching on clear surfaces (D’Anna et 
al., 2000; Moura et al., 2004) such ARs or molluscan 
shells (Fig. 8). Their dominance at the early stage of 
succession seems dependent on the timing of immer-
sion of the AR. In fact, according to a recent research 
serpulids were dominant on plates placed in summer 
and autumn, while they were overweighed by bryozo-
ans on plates placed in winter and spring (Fortič et al., 
2021). About bivalves the settlement of F. glaber was 
above discussed, whereas the employment of artificial 
substrates can drive the production of the oyster O. 
edulis and the blue mussel Mytilus galloprovincialis 
(Fabi et al., 1989; Ardizzone at al., 1989). Oysters re-
sulted very abundant on the Lignano AR and reached 
a commercial size (>70 gr) after 14 months from the 
deployment of structures (Arpa FVG, 2007), a shorter 
time compared to other areas in the Gulf of Trieste 
where about 23 months are needed (Orel & Zamboni, 
2003). The following project Ecosea used the same 
sets of AR modules for further experiment on oyster 
culture. The blue mussel did not show any dominance 
stage on modules as observed by Nicoletti et al. (2007) 
on the Fregene AR (Tyrrhenian Sea), although it was 
among characterizing species in 2007 and on PE nets 
in 2009-2011. Mytilus pediveligers are more abundant 
in the surface layer (Dobretsov & Miron, 2001) and 
the settlement on the AR could be induced by mussels 
attached on the ropes, used to link the signal buoys 
floating on the sea surface. Considering the sponge 
C. crambe, this species is normally settled on shells 
of bivalves, e.g. Arca noae, A. opercularis, M. varia, 
and gastropods such Hexaplex trunculus (Corriero et 
al., 1991) and it seems possible to farm this species 
for sponge culture purposes by means of artificial 
substrates (Padiglia et al., 2018).
The colonial ascidian B. schlosseri can be an im-
portant driver of benthic community composition, but 
it strictly depends on local environmental conditions 
(Sams & Keough, 2012). In the present study it was 
very frequent in 2007 on all AR modules, whereas it 
was almost absent in the following years. The sponges 
D. fragilis and H. mediterranea, which were constantly 
recorded since 2009, are considered respectively as 
common and characteristic in coralligenous habitat 
(sensu Ballesteros, 2006) and were constantly found 
in assemblages on the rocky outcrops in the Gulf of 
Trieste (Bettoso et al., 2023).
Based on this 5-year non-destructive monitoring 
on the AR realized thank to Project ADRI.BLU (2006) 
it is possible to conclude that the selected site was 
suitable for epibenthic settlement and in particular 
for bivalves captation. Nevertheless, according to 
Taormina et al. (2020) a 5-year monitoring is too short 
to speculate about stability or homeostasis of the 
AR in term of ecological succession. On regard the 
modules, polyethylene panel nets (PE) were functional 
only if employed during bivalve settlement, after this 
period they should be removed and cleaned for future 
employment. Considering long term settlement and 
evolution of epibenthic assemblage, TR seemed to 
give the best performance in term of species richness 
if compared to PE and EM, but a longer survey is 
needed to discriminate between long-term ecological 
successions and shorter-term variability. Anyway, on 
our opinion the use of EM and PE should be limited, 
being made by plastic material. 
ACKNOWLEDGEMENTS
The Artificial Reef in Lignano Sabbiadoro was real-
ized thank to the Project ADRI.BLU Interreg IIIA North 
Adriatic and the monitoring was funded by Regione 
Autonoma Friuli Venezia Giulia – Direzione centrale 
risorse agricole, naturali, forestali e montagna. Partic-
ular thanks are due to Giorgio Mattassi, Pietro Rossin 
and Walter de Walderstein. The present work was 
possible also thanks to the financial support from the 
Slovenian Research Agency (research core funding No. 
P1-0237).
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EPIBENTOŠKA MAKROFAVNA NA UMETNEM PODVODNEM GREBENU V SEVERNEM 
JADRANU: PET LETNI FOTOGRAFSKI MONITORING
Nicola BETTOSO, Lisa FARESI & Ida Floriana ALEFFI
Agenzia Regionale per la Protezione dell’Ambiente del Friuli Venezia Giulia (ARPA FVG), via Cairoli 14 – 33053 Palmanova (UD), Italy
e-mail: nicola.bettoso@arpa.fvg.it
Valentina PITACCO
Marine Biology Station Piran, National Institute of Biology, Fornače 41, 6330 Piran, Slovenia
POVZETEK
Umetni podvodni grebeni (UPG) so umetne strukture, ki jih uporabljajo z namenom izboljšanja ribištva 
in povečanje naravne produkcije bioloških virov. Leta 2006 so potopili tri tipe UPG blizu podvodnega 
iztoka kanalizacije. Cilji projekta so bili: (a) uporaba UPG za repopulacijo nekaterih komercialnih tarčnih 
vrst in (b) za promocijo biodiverzitete na izbranem območju. V petletnem obdobju (2007-2011) so z 
nedestruktivnimi fotografskimi metodami letno spremljali epibentoško makrofavno. Določili so 88 taksonov 
iz 8 debel, med katerimi so prevladovale spužve, mehkužci in kozolnjaki. Med različnimi tipi UPG so bile 
polietilenske panelne mreže funkcionalne za naseljevanje školjk v prvem letu, medtem ko so se betonske 
strukture izkazale za najboljše v smislu vrstne pestrosti biodiverzitete v dolgoročnem obdobju. Kakorkoli 
že, petletno obdobje je bilo prekratko za razumevanje stabilnosti oziroma homeostaze združb, ki so se v 
smislu ekološke sukcesije naselile na UPG.
Ključne besede: umetni podvodni grebeni, makrozoobentos, Jadransko morje, fotografski monitoring
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