Anali za istrske in mediteranske študije
Annali di Studi istriani e mediterranei
Annals for Istrian and Mediterranean Studies
Series Historia Naturalis, 31, 2021, 2
UDK 5                       Annales, Ser. hist. nat., 31, 2021, 2, pp. 167-326, Koper 2021  ISSN 1408-533X
KOPER 2021
Anali za istrske in mediteranske študije
Annali di Studi istriani e mediterranei
Annals for Istrian and Mediterranean Studies
Series Historia Naturalis, 31, 2021, 2
UDK 5                    ISSN 1408-533X 
                e-ISSN 2591-1783
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies
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e-ISSN 2591-1783
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ANNALES · Ser. hist. nat. · 31 · 2021 · 2
Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies
UDK 5                                                      Letnik 31, Koper 2021, številka 2                                      ISSN 1408-533X
e-ISSN 2591-1783 
VSEBINA / INDICE GENERALE / CONTENTS 2021(2)
BIOINVAZIJA
BIOINVASIONE 
BIOINVASION
Cemal TURAN, Mevlüt GÜRLEK, 
Deniz ERGÜDEN & Hakan KABASAKAL
A New Record for the Shark Fauna 
of the Mediterranean Sea: Whale shark, 
Rhincodon typus (Orectolobiformes: 
Rhincodontidae) .......................................................
Nova vrsta v favni morskih psov 
Sredozemskega morja: morski pes 
kitovec, Rhincodon typus 
(Orectolobiformes: Rhincodontidae)
Andrea LOMBARDO & Giuliana MARLETTA
New Evidence of the Ongoing 
Expansion of Okenia picoensis 
Paz-Sedano, Ortigosa & Pola, 
2017 (Gastropoda: Nudibranchia) in 
the Central-Eastern Mediterranean .........................
Novi podatki o širjenju areala vrste Okenia 
picoensis Paz-Sedano, Ortigosa & Pola, 
2017 (Gastropoda: Nudibranchia) v 
srednjem vzhodnem Sredozemskem morju
SREDOZEMSKI MORSKI PSI 
SQUALI MEDITERRANEI
MEDITERRANEAN SHARKS
Hakan KABASAKAL
A Review of Shark Biodiversity in 
Turkish Waters: Updated Inventory, 
New Arrivals, Questionable Species, 
and Conservation Issues .......................................
Pregled pestrosti morskih psov v 
turških morjih: dopolnjen seznam, 
novi prišleki, vprašljive vrste in 
naravovarstveni problemi
Hakan KABASAKAL & Erdi BAYRI
Great White Sharks, Carcharodon 
carcharias, Hidden in the Past: 
Three Unpublished Records of the 
Species from Turkish Waters ................................
Trije neobjavljeni primeri pojavljanja belega 
morskega volka, Carcharodon carcharias, 
iz turških voda izbrskani iz preteklosti
IHTIOLOGIJA
ITTIOLOGIA
ICHTHYOLOGY
Malek ALI, Vienna HAMMOUD, 
Ola FANDI & Christian CAPAPÉ 
First Substantiated Record of 
Crested Oarfish Lophotus lacepede 
(Osteichthyes: Lophotidae) from the 
Syrian Coast (Eastern Mediterranean Sea) ............
Prvi utemeljeni zapis o pojavljanju 
čopovke Lophotus lacepede 
(Osteichthyes: Lophotidae) ob 
sirski obali (vzhodno Sredozemsko morje)
Mohamed Mourad BEN AMOR, 
Khadija OUNIFI-BEN AMOR, 
Marouène BDIOUI & Christian CAPAPÉ
The Second Record of Oilfish, 
Ruvettus pretiosus (Gempylidae), 
in Tunisian Waters (Central 
Mediterranean Sea) .............................................
Drugi zapis o pojavljanju vrste 
Ruvettus pretiosus (Gempylidae) 
v tunizijskih vodah (osrednje 
Sredozemsko morje)
Okan AKYOL & Vahdet ÜNAL
On the Occurrence of Seriola fasciata 
(Carangidae) in the Eastern 
Mediterranean Sea ...............................................
O pojavljanju vrste Seriola fasciata 
(Carangidae) v vzhodnem 
Sredozemskem morju
Nassima EL OMRANI, 
Hammou EL HABOUZ, 
Abdelbasset BEN-BANI, 
Abdellatif MOUKRIM, 
Roger FLOWER & Abdellah BOUHAIMI
Age and Growth of the Pouting 
Trisopterus luscus (Linnaeus, 1758) 
(Pisces, Gadidae) from Moroccan 
Central Atlantic Waters.........................................
Rast in starost francoskega moliča 
Trisopterus luscus (Linnaeus, 1758) 
(Pisces, Gadidae) v atlantskih 
vodah osrednjega Maroka
167
173
195
217
211
181
223
205
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
Mourad CHÉRIF, Rimel BENMESSAOUD & 
Christian CAPAPÉ
Age and Growth Parameters of the Red 
Mullet Mullus barbatus (Mullidae) from 
Northern Tunisia (Central Mediterranean Sea) .........
Starostni in rastni parametri pri navadnem 
bradaču Mullus barbatus (Mullidae) iz 
severne Tunizije (osrednje Sredozemsko morje)
Yana SOLIMAN, Adib SAAD, 
Vienna HAMMOUD & Christian CAPAPÉ
Heavy Metal Concentrations in Tissues of
 Red Mullet, Mullus barbatus (Mullidae) from 
the Syrian Coast (Eastern Mediterranean Sea) ...........
Vsebnost težkih kovin v tkivih bradača, 
Mullus barbatus (Mullidae) iz sirske obale 
(vzhodno Sredozemsko morje)
Christian CAPAPÉ, Youssouph DIATTA, 
Almamy DIABY, Sihem RAFRAFI-NOUIRA & 
Christian REYNAUD
Record of a Single Clasper Specimen in 
Zanobatus schoenleinii (Chondrichthyes: 
Zanobatidae) from the Coast of Senegal 
(eastern tropical Atlantic) ..........................................
Najdba primerka vrste Zanobatus 
schoenleinii (Chondrichthyes: 
Zanobatidae) le z enim klasperjem iz 
senegalske obale (vzhodni tropski Atlantik)
FAVNA
FAVNA
FAVNA
Ana FORTIČ, Domen TRKOV, 
Lovrenc LIPEJ, Marco FANTIN & 
Saul CIRIACO
New Evidence of the Occurrence of 
Knoutsodonta pictoni (Nudibranchia, 
Onchidorididae) in the Northern Adriatic .................
Novi podatki o pojavljanju vrste 
Knoutsodonta pictoni (Nudibranchia, 
Onchidorididae) v severnem Jadranu
Noureddine BENABDELLAH, Djillali BOURAS, 
Mohammed RAMDANI & Nicolas STURARO 
Biodiversity and Structural Organization 
of Mollusk Communities in the Midlittoral 
Coastal Area Between Bouzedjar and 
Arzew (Western Algeria) ...........................................
Biodiverziteta in struktura združbe 
mehkužcev v bibavičnem območju 
med predeloma Bouzedjar in Arzew 
(zahodna Alžirija)
Rudi VEROVNIK, Nejc RABUZA, 
Miroslav REPAR, Matjaž ZADRGAL & 
Paul TOUT
On the Presence of Two-Tailed Pasha 
(Charaxes jasius (Linnaeus, 1767), 
Papilionoidea: Nymphalidae) in the 
Northeastern Adriatic Region ....................................
O pojavljanju dvorepega paše 
(Charaxes jasius (Linnaeus, 1767), 
Papilionoidea: Nymphalidae) na 
območju severovzhodnega Jadrana
Viktor BARANOV & Borut MAVRIČ
New Records of Non-Biting Midges 
(Diptera, Chironomidae) from Marine 
and Coastal Habitats of the Slovenian 
Part of the Adriatic Sea .............................................
Nove najdbe trzač (Diptera, 
Chironomidae) iz morskih in obmorskih 
habitatov v slovenskem delu Jadrana
FLORA
FLORA
FLORA
Amelio PEZZETTA, Marco PAOLUCCI & 
Mario PELLEGRINI
Le Orchidaceae del sito di interesse 
comunitario “Monte Pallano e Lecceta d’Isca 
d’Archi” e delle zone limitrofe .................................
Kukavičevke območja, pomembnega za 
skupnost “Monte Pallano e Lecceta d’Isca 
d’Archi” in sosednjih območij
DELO NAŠIH ZAVODOV IN DRUŠTEV
ATTIVITÀ DEI NOSTRI ISTITUTI E SOCIETÀ
ACTIVITIES BY OUR INSTITUTIONS AND 
ASSOCIATIONS
Marina DERMASTIA, Tina ELERŠEK, 
Jadranka JEZERŠEK, Lučka KAJFEŽ BOGATAJ, 
Matjaž KUNTNER, Tamara LAH TURNŠEK, 
Matjaž LIČER, Lovrenc LIPEJ, Miha MIKELJ, 
Izidor OSTAN OŽBOLT, Maja RAVNIKAR, 
Katja SINUR, Darja STANIČ, 
Timotej TURK DERMASTIA, Al VREZEC
Okoljski manifest ......................................................
IN MEMORIAM
Jadran FAGANELI 
V spomin prof. dr. Jožetu Štirnu (1934-2021) ...........
Kazalo k slikam na ovitku ........................................
Index to images on the cover ....................................
321
326
326
291
235
243
261
301
285
267 
251
315
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
261
received: 2021-11-17                 DOI 10.19233/ASHN.2021.31
NEW EVIDENCE OF THE OCCURRENCE OF KNOUTSODONTA PICTONI 
(NUDIBRANCHIA, ONCHIDORIDIDAE) IN THE NORTHERN ADRIATIC
Ana FORTIČ, Domen TRKOV & Lovrenc LIPEJ
Marine Biology Station Piran, National Institute of Biology, Slovenia
e-mail: Ana.Fortic@nib.si
Marco FANTIN
Sistiana Diving, Italy
Saul CIRIACO
WWF Miramare MPA, Italy
ABSTRACT
The authors present new data on the recently described nudibranch Knoutsodonta pictoni Furfaro & 
Trainito, 2017 in the Gulf of Trieste (northern Adriatic Sea). We hereby present the first record of this 
species in Slovenian territorial waters. A total of 22 specimens of K. pictoni were recorded by diving in the 
period from 2017 to 2021 at five localities in the Gulf of Trieste. All specimens were found in precorallig-
enous habitats in the depth range of 5 to 9 m, feeding on the encrusting bryozoan Reptadeonella violacea 
(Johnston, 1847). Numerous finds indicate that this only recently described and therefore little known 
nudibranch is much less rare than previously thought.
Key words: Knoutsodonta pictoni, Gulf of Trieste, sea slug, precoralligenous, cryptic species
NUOVE PROVE DELLA PRESENZA DI KNOUTSODONTA PICTONI (NUDIBRANCHIA, 
ONCHIDORIDIDAE) NELL’ADRIATICO SETTENTRIONALE
SINTESI
Gli autori presentano nuovi dati su una specie di nudibranco descritta recentemente, Knoutsodonta 
pictoni Furfaro & Trainito, 2017, nel Golfo di Trieste (Adriatico settentrionale). L’articolo fornisce il primo 
ritrovamento della specie nelle acque territoriali slovene. Un totale di 22 esemplari di K. pictoni sono 
stati trovati durante le immersioni effettuate nel periodo dal 2017 al 2021, in cinque località del Golfo di 
Trieste. Tutti gli esemplari sono stati rinvenuti in habitat precoralligeni, nell’intervallo di profondità da 5 
a 9 m, mentre si nutrivano del briozoo incrostante Reptadeonella violacea (Johnston, 1847). I numerosi 
ritrovamenti indicano che questo nudibranco, descritto solo recentemente e quindi poco conosciuto, è 
molto meno raro di quanto si pensasse.
Parole chiave: Knoutsodonta pictoni, Golfo di Trieste, lumaca di mare, precoralligeno, specie criptica
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
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Ana FORTIČ et al.: NEW EVIDENCE OF THE OCCURRENCE OF KNOUTSODONTA PICTONI (NUDIBRANCHIA, ONCHIDORIDIDAE) IN THE NORTHERN ..., 261–266
INTRODUCTION
Knoutsodonta pictoni Furfaro & Trainito, 2017 
(Fig. 1) is a recently described nudibranch of the fam-
ily Onchidorididae Gray, 1827. It is distributed in the 
western Mediterranean Sea in Spain (Ballesteros et al., 
2016; OPK Opistobranquis, 2020), in the Tyrrhenian 
and Ligurian Seas in Italy (Betti et al., 2017; Furfaro & 
Trainito, 2017), and in the northeast Atlantic Ocean in 
Ireland and Scotland (Hallas & Gosliner, 2015; Furfaro 
& Trainito, 2017 and references therein; nudibranch.org, 
2021). The first record of the species in the Adriatic Sea 
was reported by Furfaro & Trainito (2017) in Sistiana, 
Gulf of Trieste. Morphological features characteristic of 
the species are: elliptical and dorsally flattened body with 
bristly appearance; base colour dark brown with blue and 
white speckles on the mantle; rhinophores white with 9 
to 11 lamellae; gills dark brown with 9 to 10 bipinnate 
branchial leaves (Furfaro & Trainito, 2017).
The aim of this short note is to provide new data on 
the occurrence of this species in the Gulf of Trieste and 
on the ecology of this nudibranch, as well as reporting the 
first record of this species for the Slovenian coasts.
MATERIAL AND METHODS
Observations and sampling were carried out by 
scuba diving in the Italian and Slovenian parts of the 
Gulf of Trieste between 2017 and 2021, more specifi-
cally in Sistiana and Sistiana Castelreggio in the north-
ern part of the Gulf of Trieste, in Barcola near the town 
of Trieste, and in the southern part of the gulf, Piranček 
and Fiesa near the town of Portorož.
In Italy, the nudibranchs were observed from 2017 to 
2020. Specimens were photographed with a Sea & Sea 
2G camera with a Sea & Sea DS1 strobe using Nauticam 
CMC1 macro lens, and identified in situ. Depth and tem-
perature were recorded at all sites. 
In Slovenia, several bryozoan species were collected 
during a marine biodiversity survey in Natura 2000 sites 
in summer 2021. Depth and temperature were recorded. 
Nudibranchs and the bryozoan colonies on which they 
were found were identified in the laboratory and photo-
graphed under a stereomicroscope (Olympus SZX16) with 
a digital camera (Olympus DP25), with the nudibranchs 
subsequently deposited in the malacological collection. 
Specimens were identified using the diagnostic fea-
tures described by Furfaro & Trainito (2017). Systematics 
and validity of names were checked using the World 
Register of Marine Species [WoRMS].
RESULTS AND DISCUSSION
A total of 22 specimens (20 of which found in Italy 
and 2 in Slovenia) and 6 egg masses (found only in Italy) 
were photographed over a five-year period (from 2017 
to 2021; see Tab. 1 for details). All specimens displayed 
the same colour pattern with little chromatic variability. 
In general, this species was found in winter and summer 
on precoralligenous rocky bottoms 5 to 9 m deep and in 
a temperature zone ranging from 7 to 26°C. The habitats 
where the specimens were observed were located under 
a b c
Fig. 1: Knoutsodonta pictoni; a – specimen feeding on the bryozoan Reptadeonella violacea, b – two speci-
mens, and c – close-up of a specimen with visible white lamellate rhinophores (Photo: M. Fantin).
Sl. 1: Knoutsodonta pictoni; a – primerek se prehranjuje na mahovnjaku Reptadeonella violacea, b – dva 
primerka, in c – bližinski posnetek primerka z vidnima lamelastima rinoforjema (Foto: M. Fantin).
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
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Fig. 2: Map of the Gulf of Trieste with five localities where the specimens of Knoutsodonta pictoni were found 
(black dots). The numbers next to the black dots represent the localities listed in Tab. 1.
Sl. 2: Zemljevid Tržaškega zaliva s petimi lokalitetami, kjer so bili najdeni primerki vrste Knoutsodonta pictoni 
(črni krogci). Številke poleg krogcev označujejo lokalitete, ki so navedene v Tab. 1.
date
locality 
number
locality GPS N GPS E n
egg 
mass
depth 
(m)
T (°C)
14.1.2017 1 Sistiana 45°46’7.88’’ 13°37’21.19” 2 1 7 7
21.1.2017 2 Barcola 45°41’30.53’’ 13°44’20.26” 2 1 4 7
29.1.2017 1 Sistiana 45°46’7.88” 13°37’21.19’’ 2  8 8
30.1.2017 3 Sistiana Castelreggio 45°45’58.21” 13°38’2.04” 2  8 8
4.2.2017 3 Sistiana Castelreggio 45°45’58.21” 13°38’2.04” 2 1 9 9
5.2.2017 3 Sistiana Castelreggio 45°45’58.21” 13°38’2.04” 2  8 9
18.2.2017 1 Sistiana 45°46’7.88” 13°37’21.19’’ 2 1 9 9
10.2.2018 3 Sistiana Castelreggio 45°45’58.21” 13°38’2.04” 2 1 7 10
24.2.2018 3 Sistiana Castelreggio 45°45’58.21” 13°38’2.04” 2 1 8 10
10.3.2018 1 Sistiana 45°46’7.88” 13°37’21.19’’ 1  9 11
2.6.2020 1 Sistiana 45°46’7.88” 13°37’21.19’’ 1  8,5 16
20.7.2021 4 Fiesa 45°31’38.4’’ 13°34’42.54’’ 1  7 21
29.7.2021 5 Piranček 45°31’16.75’’ 13°33’57.45’’ 1  5 26
Tab. 1: Data on the occurrence of Knoutsodonta pictoni in the Gulf of Trieste in the period 2017‒2021. 
Tab. 1: Podatki o pojavljanju vrste Knoutsodonta pictoni v Tržaškem zalivu v obdobju med 2017 in 2021.
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stones, on molluscan shells, and on rocks. In Sistiana 
the nudibranch was regularly observed in 2017, 2018, 
and 2020, in Sistiana Castelreggio it was recorded in the 
winters of 2017 and 2018, and in Barcola only once, 
in winter 2017. The spawning activities were observed 
from late January to May. The egg masses were white, 
concentrically folded and ribbon-shaped (Fig. 3a). On 
one occasion, an oviphagous nudibranch, Favorinus 
branchialis (Rathke, 1806), was photographed devouring 
an egg mass of K. pictoni (Fig. 3b). With regard to find-
ings along the Slovenian coast, only two specimens were 
found in summer 2021, both small-sized (between 3 and 
4 mm), probably juveniles.
The ecology of K. pictoni, being this nudibranch a 
recently described species, is poorly known. However, 
the numerous findings reported in this study indicate 
that it is far less rare than previously thought. All 
the specimens collected were found feeding on the 
encrusting cheilostomatid bryozoan Reptadeonella 
violacea (Johnston, 1847). The same was observed 
by Furfaro & Trainito (2017), suggesting a preferential 
predatory relationship. In fact, it is not uncommon for 
nudibranchs to feed on one prey species only (Todd & 
Havenhand, 1989). R. violacea is a common bryozoan 
species in the shallow waters of the Mediterranean 
Sea, living as an epiphyte in Posidonia meadows, on 
algae (Novosel, 2005), on the underside of stones, and 
on molluscan shells. Nudibranchs are mostly known 
as vividly coloured heterobranchs, while species with 
cryptic colouration are less well known. They use 
various camouflage strategies such as homochromy, 
countershading, and cryptic or disruptive colouration 
(Todd, 1981). These cryptic species, such as K. pictoni, 
were often overlooked in the past, due to their excel-
lent camouflage. In fact, to date, only three species of 
the family Onchidorididae [K. pictoni; K. neapolitana 
(Delle Chiaje, 1841) and K. depressa (Alder & Han-
cock, 1842)] have been recorded in the Gulf of Trieste 
and throughout the northern Adriatic Sea (Ciriaco 
& Poloniato, 2016; Zenetos et al., 2016; Furfaro & 
Trainito, 2017; Lipej et al., 2018; present study). There-
fore, continuous monitoring is necessary to extend our 
knowledge of these particular species, which often go 
unnoticed.
ACKNOWLEDGMENTS
 
The authors would like to dedicate this paper to 
Barbara Camassa, a passionate diver and underwater 
photographer. Our special thanks go to Borut Mavrič, 
who provided us with samples of bryozoans for analy-
sis, and Milijan Šiško, who drew the map of the locali-
ties. Finally, we would like to thank two anonymous 
reviewers who helped us improve the article. 
a b
Fig. 3: Spawn of Knoutsodonta pictoni; a – white ribbon with eggs on the bryo-
zoan Reptadeonella violacea, b – oviphagous nudibranch Favorinus branchialis 
feeding with the eggs of K. pictoni (Photo: M. Fantin). 
Sl. 3: Mrest vrste Knoutsodonta pictoni; a – beli svitek z jajci na mahovnjaku vrste 
Reptadeonella violacea, b – ovifagni gološkrgar vrste Favorinus branchialis se 
hrani z jajci vrste K. pictoni (Foto: M. Fantin). 
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NOVI PODATKI O POJAVLJANJU VRSTE KNOUTSODONTA PICTONI (NUDIBRANCHIA, 
ONCHIDORIDIDAE) V SEVERNEM JADRANU
Ana FORTIČ, Domen TRKOV & Lovrenc LIPEJ
Marine Biology Station Piran, National Institute of Biology, Slovenia
e-mail: Ana.Fortic@nib.si
Marco FANTIN
Sistiana Diving, Italy
Saul CIRIACO
WWF Miramare MPA, Italy
POVZETEK
Avtorji poročajo o novih najdbah pred kratkim opisanega polža gološkrgarja vrste Knoutsodonta pictoni 
Furfaro & Trainito, 2017 v Tržaškem zalivu (severni Jadran). Navajajo prvi zapis o pojavljanju te vrste za 
slovenske vode. V obdobju med 2017 in 2021 je bilo na potapljaških vzorčenjih  popisanih skupno 22 pri-
merkov vrste K. pictoni na petih lokalitetah v Tržaškem zalivu. Vsi primerki so bili najdeni v prekoraligenu 
v globinskem pasu med 5 in 9 m, kjer so se prehranjevali s skorjastim mahovnjakom vrste Reptadeonella 
violacea (Johnston, 1847). Številne najdbe kažejo, da je ta pred kratkim opisana in slabo poznana vrsta 
pogostejša, kot so domnevali doslej. 
Ključne besede: Knoutsodonta pictoni, Tržaški zaliv, gološkrgarji, prekoraligen, kriptična vrsta 
ANNALES · Ser. hist. nat. · 31 · 2021 · 2
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