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ANNALES · Ser. hist. nat. · 28 · 2018 · 1
short scientifi c article                 DOI 10.19233/ASHN.2018.03
received: 2018-03-05
ON THE RARE AND LESS KNOWN SHAMEFACED CRAB 
CALAPPA GRANULATA (BRACHYURA, CALAPPIDAE) 
IN THE NORTHERN ADRIATIC SEA
Nicola BETTOSO
ARPA FVG, via A. La Marmora 13, 34139 Trieste, Italy
e-mail: nicola.bettoso@arpa.fvg.it
Marin KIRINČIĆ
Natural History Museum, Lorenzov prolaz 1, 51000 Rijeka, Croatia
Borut MAVRIČ & Lovrenc LIPEJ
Marine Biology Station, National Institute of Biology, Fornače 41, SI-6330 Piran, Slovenia
ABSTRACT
On 1st August 2016 and 23th July 2017 two specimens of the shamefaced crab Calappa granulata (Linnaeus, 
1758) were caught in the northernmost area of the Adriatic Sea and represent the second and third offi cial record, 
respectively. More recently the records of this commercial species start to be no more unusual in this northern region. 
It is too early to express any reliable comment regarding whether or not this species established a breeding popu-
lation in this area, however we could consider its northward extension as another consequence of global warming.
Key words: shamefaced crab, Calappa granulata, northern Adriatic, global warming
IL RARO E POCO CONOSCIUTO GRANCHIO MELOGRANO CALAPPA GRANULATA 
(BRACHYURA, CALAPPIDAE) NELL’ADRIATICO SETTENTRIONALE
SINTESI
Il primo agosto 2016 e il 23 luglio 2017 sono stati catturati due esemplari di granchio melograno Calappa 
granulata (Linnaeus, 1758) nell’area più settentrionale del mare Adriatico e ne rappresentano rispettivamente la 
seconda e la terza segnalazione uffi ciale. Attualmente le catture di questa specie commerciale non sembrano più 
inconsuete come nel passato. È ancora troppo presto per avanzare ipotesi riguardo al suo insediamento in pianta 
stabile su quest’area, tuttavia l’espansione verso nord di questa specie può costituire un ulteriore conseguenza del 
riscaldamento globale dei mari.
Parole chiave: granchio melograno, Calappa granulata, Adriatico settentrionale, riscaldamento globale
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ANNALES · Ser. hist. nat. · 28 · 2018 · 1
Nicola BETTOSO et al.: ON THE RARE AND LESS KNOWN SHAMEFACED CRAB CALAPPA GRANULATA (BRACHYURA, ..., 15–22
INTRODUCTION
The shamefaced crab Calappa granulata (Linnaeus, 
1758) is a sublittoral species known from the Mediter-
ranean Sea and adjacent Atlantic Ocean from Portugal 
to Mauritania, including the Azores, Madeira, the 
Canary Islands and the Cape Verde Islands (Manning & 
Holthuis, 1981; Števčić, 1990). More recent records in 
the Mediterranean Sea are the Gulf of Taranto (Ionian 
Sea) (Pastore, 1995), the Strait of Sicily (Spanò et al., 
2004), the coastal waters of the Sea of Marmara (Artűz, 
2006) and Edremit Bay (Aegean Sea) (Balkis & Kurun, 
2008). In the Adriatic Sea, Stossich (1880) mentioned 
this species as rare in Split, Hvar, Vis and Korčula. After 
110 years Števčić (1990) confi rmed the rarity of this crab 
in the Middle and South Adriatic where it was recorded 
in waters off Šibenik, Split, Dubrovnik, Sestrunje, Hvar, 
Korčula and Vis. The last records from the southern 
Adriatic arises to Ungaro et al. (2005) who found some 
individuals during trawling operations in the framework 
of the E.U. Project MEDITS, moreover Marković et al. 
(2017) recorded this species also in Montenegron waters 
in Kotor Bay and Herceg Novi Bay. On December 2010, 
a specimen was caught in the northern Adriatic Sea 
northwest off Umag, on the western coast of the Istrian 
Peninsula, a region substantially further North than 
any previous records in the Adriatic (Dulčić & Tutman, 
2012). In 2016 and 2017 another two specimens were 
collected off Istrian Peninsula and more recently the 
records of this commercial species start to be no more 
unusual in this northern region.
MATERIAL AND METHODS
On 23th July 2017, a specimen of Calappa granulata 
was caught 4 Nm [nautical miles: approx. 7.4 Km] off 
Funtana by a trammel net at about 20 m depth on a 
muddy sand bottom. Previously another specimen was 
caught on 1st August 2016 in bottom-set gillnet 2 Nm 
[nautical miles: approx. 3.7 Km] off Piran (Fig. 1).
Specimens were identifi ed following Holthuis (1987) 
and compared to the two other deposited in the Natural 
History Museum of Rijeka (Catalogue numbers: C195 and 
C1540). C195 was deposited on January 1996 and col-
lected in the waters near Cres Island, whereas C1540 was 
sampled in July 2012 from Dubrovnik; the latter caught 
by trawling at 200 m depth (Fig. 1). Four investigated 
specimens were sexed and the following morphometric 
measurements were performed: Rostrum Width (RW), 
Carapace Width (CW), Carapace Length (CL), Posterior 
Margin Width (PMW), Abdomen Length (AL), Abdomen 
Width (AW), Dactilo-claws Length (DL), Propodium 
Length (PL), Cheliped Length (ChL), Cheliped Height 
(ChH), Cheliped Width (ChW) and 1st male Gonopod 
Length (G1L). Isomorphism between specimens was 
tested by Pearson linear correlation coeffi cient (PCC).
RESULTS AND DISCUSSION
Morphometric measurements of Calappa granulata 
are showed in Tab. 1 and Pearson linear correlation 
coeffi cient revealed a complete isomorphism between 
specimens (Tab. 2). All specimens were adult males, as 
well as that recorded by Dulčić & Tutman (2012). The 
species can reach 8 cm CL and 11 cm CW (Coudre et 
al., 2013). In the Mediterranean, it lives on sandy mud 
and muddy detritus at depths between 13 and 400-700 
m (Manning & Holthuis, 1981; Abelló et al., 1988). The 
specimens from Piran and Funtana (Fig. 2) represent 
the second and third offi cial records respectively in the 
northernmost area of the Adriatic Sea. Among oldest 
fi shermen in the Istrian peninsula, they do not remem-
ber any such crabs in the past. Until now the species 
was not recorded in the checklists of decapod fauna in 
Gulf of Trieste (sensu Manning & Števčić, 1982) and in 
the Rijeka Bay (sensu Zavodnik & Kovačić, 2000).
Fig. 1: Records of Calappa granulata in the Adriatic 
Sea: Piran 2016 and Funtana 2017 (the present work), 
Umag 2010 (Dulčić & Tutman, 2012), Cres 1996 and 
Dubrovnik 2012 are referred to specimens deposited in 
the Natural History Museum of Rijeka; the circle shows 
the geographic area indicated by Stossich (1880) and 
Števčić (1990).
Sl. 1: Podatki o pojavljanju vrste Calappa granulata v 
Jadranskem morju: Piran 2016 in Funtana 2017 (priču-
joče delo), Umag 2010 (Dulčić & Tutman, 2012), Cres 
1996 in Dubrovnik 2012 pa se nanašajo na primerke v 
zbirki Prirodoslovnega muzeja na Reki; krogec označu-
je območje, ki ga navajata Stossich (1880) in Števčić 
(1990).
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ANNALES · Ser. hist. nat. · 28 · 2018 · 1
Nicola BETTOSO et al.: ON THE RARE AND LESS KNOWN SHAMEFACED CRAB CALAPPA GRANULATA (BRACHYURA, ..., 15–22
It is too early to express any reliable comment regard-
ing whether or not this species established a breeding 
population in the area as suggested by Dulčić & Tutman 
(2012). The same authors hypothesized that pelagic 
larvae of the shamefaced crab could be transported by 
currents, since the hydrodynamic characteristics of the 
Adriatic Sea support a hypothesis of passive transport. 
These records could be also explained by the effect of 
the Ionian water ingressions in the Adriatic (Dulčić & 
Grbec, 2000) and further support the decadal variability 
of water mass properties of the basin (Civitarese et al., 
2010). Nevertheless C. granulata shows a preference for 
tropical waters (Spanò et al., 2004) and its northward 
extension could be another consequence of global 
warming. Thanks to palaeontological records, we know 
that C. granulata populated the Pliocene sea in corre-
spondence to the present Po Plain (Garassino & Pasini, 
2013; Pasini & Garassino, 2013), which was a wide and 
deep gulf much warmer than today (Marchetti et al., 
2017). Climate change and the impacts of commercial 
fi shing are shifting the benthic community structure, 
in particular bottom trawling has caused a widespread 
decline of traditional exploited stocks (Bastari et al., 
2017) being replaced by newcomer, although shipping 
as a potential vector of arrival cannot be excluded 
(Dulčić & Tutman, 2012). In the trawling ground of 
the meso-Adriatic depression (Pomo pit), in fact, the 
previously dominant squat lobster Munida intermedia 
has been totally replaced by the newcomer M. rutllanti, 
the latter being fi rst recorded in 2003 for Italian seas. 
In the same period the shrimp Parapenaeus longirostris 
became an important fi sheries resource, while the 
traditional Norway lobster stock (Nephrops norvegicus) 
dramatically decreased, partly due to a long-lasting 
over-fi shing (Froglia, 2017).
Tab. 1: Morphometric measurements of Calappa granu-
lata (in mm).
Tab. 1: Morfometrične meritve primerkov vrste Calappa 
granulata (v mm).
Specimen C 195 C 1540 Piran Funtana
SEX M M M M
RW 10.6 10.1 10.6 9.9
CW 104.5 80.3 97.7 103.5
CL 77.7 59.8 73.7 77.5
PMW 33.9 28.4 28.5 34.4
AL 55.7 41.9 46.1 48.3
AW 21.3 15.3 19.9 20.4
DL 38.9 28.6 35.9 36.4
PL 69.9 54.1 60.8 67.2
CIL 58.1 45.4 53 58.8
CIH 59.1 43.2 54.8 54.9
CIW 18.1 13.2 17 17.9
G1L 27.3 20.6 25.7 25.9
Tab. 2: Pearson linear correlation coeffi cient (PCC).
Tab. 2: Primerjava primerkov s Pearsonovim koefi cien-
tom korelacije (PCC).
PCC C 195 C 1540 Piran
C 1540 1.00   
Piran 1.00 0.99  
Funtana 1.00 1.00 1.00
Fig. 2: Calappa granulata (Linnaeus, 1758): A specimen 
from Piran (CL♂ = 73.7 mm, CW♂ = 97.7 mm, Wt = 186 
g). B specimen from Funtana (CL♂ = 77.5 mm, CW♂ = 
103.5 mm, Wt = 195.7 g).
Sl. 2: Calappa granulata (Linnaeus, 1758): A - Primerek 
iz Pirana (CL♂ = 73,7 mm, CW♂ = 97,7 mm, Wt = 186 g). 
B – primerek iz Funtane (CL♂ = 77,5 mm, CW♂ = 103,5 
mm, Wt = 195,7 g).
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ANNALES · Ser. hist. nat. · 28 · 2018 · 1
Nicola BETTOSO et al.: ON THE RARE AND LESS KNOWN SHAMEFACED CRAB CALAPPA GRANULATA (BRACHYURA, ..., 15–22
The decapod fauna of the northernmost part of the 
Adriatic was not given any particular scientifi c attention 
in the past in comparison with other Adriatic areas (Lipej 
et al., 2010). Only few works have been published such 
as the paper published by Manning & Števčić (1982) on 
the decapod fauna of Piran. Additional records of deca-
pods from the Piran area were given in older works by 
Stossich (1880), Graeffe (1902) and Pesta (1918). Most 
of the time, in fact, we get rare and/or alien fi sh and 
crabs thanks to the precious collaboration with fi sher-
men, who experience daily work at sea. In the future C. 
granulata could become a new resource, a new stable 
component of the northern Adriatic fauna and/or a new 
competitor for other decapods species. For these rea-
sons an establishment of a network of different groups 
who are somehow dealing with the biodiversity of the 
marine environment such as scientists, conservators, 
fi shermen, divers, underwater photographers and oth-
ers should be an important step towards the assessment 
of populations of C. granulata and other interesting and 
commercial newcomer species in the area (sensu Lipej 
et al., 2010).
ACKNOWLEDGEMENTS
We are grateful to Mario Ipsa, the oldest fi sherman in 
Funtana, for providing us a specimen of shamefaced crab 
and the precious information about frequent captures of 
this species in the area. Thanks are due also to Alfi o 
Perissinotti for providing photo of Funtana specimen.
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ANNALES · Ser. hist. nat. · 28 · 2018 · 1
Nicola BETTOSO et al.: ON THE RARE AND LESS KNOWN SHAMEFACED CRAB CALAPPA GRANULATA (BRACHYURA, ..., 15–22
O REDKI IN MANJ ZNANI VRSTI RAKOVICE CALAPPA GRANULATA (BRACHYURA, 
CALAPPIDAE) V SEVERNEM JADRANU
Nicola BETTOSO
ARPA FVG, via A. La Marmora 13, 34139 Trieste, Italy
e-mail: nicola.bettoso@arpa.fvg.it
Marin KIRINČIĆ
Natural History Museum, Lorenzov prolaz 1, 51000 Rijeka, Croatia
Borut MAVRIČ & Lovrenc LIPEJ
Marine Biology Station, National Institute of Biology, Fornače 41, SI-6330 Piran, Slovenia
POVZETEK
Dva primerka kalape Calappa granulata (Linnaeus, 1758) sta bila ujeta 1. avgusta 2016 in 23. julija 2017 na 
skrajnem severu Jadrana in predstavljata drugi in tretji podatek o pojavljanju te vrste v severnem Jadranu. V zadnjih 
letih so podatki o pojavljanju te komercialno pomembne rakovice v Jadranu pogostejši. Za zdaj je še prezgodaj 
ugotavljati, ali se vrsta na tem območju že razmnožuje, vsekakor pa gre za še en primer širjenja vrst proti severu 
zaradi globalnega segrevanja.
Ključne besede: kalapa, Calappa granulata, severni Jadran, globalno segrevanje
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Nicola BETTOSO et al.: ON THE RARE AND LESS KNOWN SHAMEFACED CRAB CALAPPA GRANULATA (BRACHYURA, ..., 15–22
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