Original scientific article	UDK 597.556.33:591.9(262.32)
Received: 2013-11-05
NEW NORTHERNMOST RECORD OF THE BLUNTHEAD PUFFERFISH, SPHOEROIDES PACHYGASTER (OSTEICHTHYES: TETRAODONTIDAE)
IN THE MEDITERRANEAN SEA
Lovrenc LIPEJ & Borut MAVRIČ
Marine Biology Station, National Institute of Biology, SI-6330 Piran, Fornače 41, Slovenia
E-mail: lipej@mbss.org
Dejan PALISKA
University of Ljubljana, Faculty of Maritime Studies and Transport, SI-6320 Portorož, Pot pomorščakov 4, Slovenia
ABSTRACT
A specimen of blunthead pufferfish Sphoeroides pachygaster (Müller & Troschel, 1848) was caught in waters off Piran on 22 November 2012. This record represents the first catch in Slovenian waters and the northernmost occurrence of this species in the Adriatic and the Mediterranean Sea, as well. The blunthead pufferfish began its rapid spread from the east Atlantic towards the Mediterranean in 1979 reaching its northernmost extent in the northern Adriatic Sea thirty years later.
Key words: blunthead pufferfish, Sphoeroides pachygaster, northern spread, Adriatic Sea
NUOVA SEGNALAZIONE A NORD DEL PESCE PALLA LISCIO, SPHOEROIDES PACHYGASTER (OSTEICHTHYES: TETRAODONTIDAE), NEL MARE MEDITERRANEO
SINTESI
Un esemplare di pesce palla liscio, Sphoeroides pachygaster (Müller & Troschel, 1848), è stato catturato nelle acque al largo di Pirano, il 22 novembre 2012. Si tratta della prima cattura di tale specie in mare sloveno e della sua segnalazione più settentrionale nell'Adriatico e nel Mediterraneo. Il pesce palla liscio si sta velocemente espandendo dall'Atlantico orientale al Mediterraneo dal 1979, raggiungendo la sua massima estensione a nord, nell'Adriatico settentrionale, trent'anni più tardi.
Parole chiave: pesce palla liscio, Sphoeroides pachygaster, espansione settentrionale, mare Adriatico
INTRODUCTION
Mediterranean fish fauna has faced many changes over the last decades. Many fish species were recorded for the first time in the Mediterranean Sea after the opening of the Suez Canal in 1865 while other newcomers arrived through the Gibraltar Strait. Some authors have considered the Mediterranean Sea to be one of the main hotspots of marine bioinvasion on the planet (Quignard & Tomasini, 2000).
The continuous arrival of new species has also been confirmed, although to a much lesser extent, in the Adriatic Sea (see for example Dulcic et al., 2003; Lipej & Dulcic, 2004; Dragicevic & Dulcic, 2010; Dulcic & Dragicevic, 2011). During the same period, native fish fauna experienced some changes as well. Some thermo-philous southern species extended their area of distribution to the north. The northward spread of southern species is caused by the ongoing phenomenon of global warming (Francour et al., 1994; Massuti et al., 2010).
The aim of this paper is to describe a specimen of blunthead pufferfish Sphoeroides pachygaster (Müller & Troschel, 1848) from the Gulf of Trieste in order to provide an overview of species spatial distribution in the Mediterranean and Adriatic Seas and analyze the spread of the blunthead pufferfish in both areas. The species is circumglobally distributed in the temperate and tropical Atlantic, in the Indian Ocean, in waters off Japan and the Hawaiian archipelago (Tortonese, 1986) and in the seas off Australia and New Zealand (Hardy, 1981). It is also present in the Mediterranean Sea; however the number of reports of this species is still rather low.
MATERIAL AND METHODS
The specimen of the blunthead pufferfish (Fig. 1) was caught in waters 5 nm west of Piran (45o35'43 N, 13o29'01 E) on 22 November 2012. It was caught from the muddy bottom at 22 m of depth. It was readily identified as the blunthead pufferfish (Müller & Troschel,
Fig. 1: Blunthead pufferfish (Sphoeroides pachygaster) caught in waters off Piran in November 2012 (Photo: B. Mavric).
Sl. 1: Riba napihovalka Sphoeroides pachygaster, ujeta v vodah pri Piranu novembra 2012 (Foto: B. Mavric)
1848) according to the identification key of Tortonese (1986). It was accurately measured to the nearest millimetre and weighed to the nearest gram. All measurements (according to Jardas, 1996) and meristic counts are presented in Table 1. The specimen is housed in the ichthyological collection of the Marine Biology Station (National Institute of Biology) in Piran.
In order to understand the phenomenon of the blunthead pufferfish spread to the northernmost area of the Adriatic Sea and in the Mediterranean Sea as well, all available literature records of this species in the Mediterranean Sea and the adjacent eastern Atlantic were gathered. On the basis of more than 100 records (from 69 reported sources; see Appendix I) of the blunthead pufferfish in the eastern Atlantic and in the Mediterranean Sea we produced a map of the area with all records presented (Fig. 2). In many sources catchment locations were only approximately defined e.g. Sicily channel, Susac Island etc. In those cases the coordinates have been chosen either within the centre of the area or territory or randomly near the territory. Although the data contain measurement errors in terms of coordinates, these errors are not problematic in this particular study of the wide area spread of the blunthead pufferfish. Another important consideration regarding the data set, when studying spatial distributions and their evolution over time, is that multiple records are reported at the same location in different time periods. This could mean that the species had settled down and is constantly present in the area. Using such data in the process of modelling leads to an inclusion of uncertainty in the data and produces errors in prediction location/trends (more in Gabrosek & Cressie, 2002). To deal with this problem a raster grid with cell size 0.5 degree latitude/longitude was created in ArcGis 9.3 and overlaid on sample data. If a single cell occupied more than one catchment location, the minimum value (year of the first record in the area) was assigned to the cell. For each single raster cell the centroid with minimum value as an attribute was extracted. The transformed dataset, year of first records within the 0.5 degree grid, were used in further analysis. This procedure has also positively influenced the effect of preferential sampling, since for the areas with dense concentrated samplings only one value was assigned, making a sample more evenly distributed in the whole area. A drawback of such a procedure is that the data on species spatial density were partially lost (32% of the original records were lost mostly in the area of high records density).
Our aim was to seek indices that could help us reveal the spatial pattern of the blunthead pufferfish population and its change over time. A good index for detecting change in spatial distribution over time is a distributional centroid and the variance of spatial distribution (Hollowed, 1992). The standard deviation ellipse is one of the tools incorporated in Esri ArcMap 9.3 software and has been used to demonstrate shifts in spatial
Fig. 2: Records of the blunthead pufferfish (S. pachygaster) in the Mediterranean Sea and eastern North Atlantic. The three directional distribution ellipses, calculated for three different time periods, indicate the shift in spatial distribution in time. The red square represents the first record off the European Atlantic coast. Sl. 2: Zapisi o pojavljanju ribe napihovalke S. pachygaster v vzhodnem Atlantiku in Sredozemlju. Tri elipse, izračunane za tri različna časovna obdobja, kažejo prostorsko smer širjenja v posameznih obdobjih. Rdeči kvadrat prikazuje prvi zapis ob atlantski obali Evrope.
distribution over time. Spatial autocorrelation, within the entire study area, was tested using Global Morans I statistics (Anselin, 1995). In order to get a better insight into how local variations are clustered, Getis-Ord Gi statistics were also calculated (Getis & Ord, 1992). Getis-Ord Gi statistics (hereafter called Ord Gi) indicate, for any given location, how individual location is associated with the values of surrounding locations. Ord Gi statistics calculate standardized Z scores for any single location. High negative or positive Z scores indicate statistically significant spatial clustering formed by low or high values (Getis Ord, 1992; Ord & Getis, 1995). Absolute values higher than 1.96 are statistically significant for spatial clustering at the 0.05 level. Values around zero indicate that no apparent spatial clustering exists (neighbours in the selected area have almost random values). The ESDA (Explanatory Spatial Data Analysis) was carried out using GeoDa ver.0.95 (Anselin, 1995). All the parameters that have been used to demonstrate spatial spreading were calculated either in Esri ArcGis 9.3 or in GeoDa.
RESULTS AND DISCUSSION Description and identification
All measurements and meristic counts are presented in Table 1. The studied specimen is among the biggest pufferfish caught in the Adriatic Sea to date, second only to the specimen, caught off Montenegro in 2008 (Joksi-movic & Mandic, 2008), which measured 450 mm in total length (Tab. 2). The specimen was identified by its typical stout and inflatable body with rounded snout, big head and skin without scales. In the mouth four large teeth (two in each jaw) form a beak. Eyes are rather big and oval in shape. A small dorsal fin is placed above the anal fin of similar size and shape. The caudal fin is slightly concave. The pelvic fin is absent. The body has smooth skin, without any scales, prickles or spines (as typical for the other species of this genus, recently confirmed in the Mediterranean Sea, namely Sphoeroides marmoratus). The colour of the dorsal surface and the flanks is greyish to olive green with many pale dots. The
Tab. 1: Morphometric data and meristic characters of the blunthead pufferfish specimen caught in waters off Piran in November 2012.
Tab. 1: Morfometrični in meristični podatki o primerku napihovalke, ujete novembra 2012 v vodah pri Piranu
Morphometric parameter	mm	% of Total length
Total length	348	100
Standard length	300	86.21
Head length	99	28.45
Head height	72	20.69
Head width	77	22.13
Eye horizontal diameter	23	6.61
Eye vertical diameter	16	4.60
Interorbital space	38	10.92
Snout length	49	14.08
Postorbital length	34	9.77
Width of peduncle	49	14.08
Width of gill opening	15	4.31
Predorsal length	220	63.22
Preanal length	232	66.67
Dorsal fin length	35	10.06
Dorsal fin base length	16	4.60
Anal fin length	37	10.63
Anal fin base length	15	4.31
Pectoral fin length	39	11.21
Caudal fin length	40	11.49
Body thickness	93	26.72
Body height	93	26.72
Internarial space	30	8.62
Meristic counts		
Dorsal fin rays	8	
Anal fin rays	8	
Pectoral fin rays	14	
Caudal fin rays	10	
ventral side is whitish. The meristic counts and morphometric data agree well with previously published data on the Mediterranean Sea (for example Dulcic, 2002).
Records of S. pachygaster in Mediterranean
Among the newcomers in the Mediterranean and the Adriatic Seas, the blunthead pufferfish deserved appropriate scientific attention over the last decades, beginning with its first Mediterranean appearance in 1979 in waters off the Balearic Islands (Oliver, 1981) (Fig.2 and Appendix I). After that time many records from various areas of the western Mediterranean Sea were published (see Dulcic, 2002 for site locations). This circumglobal temperate and tropical species rapidly dispersed thro-
ughout the Mediterranean Sea (Ragonese et al., 1997) and it is considered as rather common in the western basin and in certain areas of the eastern basin (Golani et al., 2002). In the eastern Mediterranean, the species was first reported by Golani (1996), who mentions a specimen collected by a trawl off Ashdod (Israel) in 1991 (Golani, 1996). In the eastern Mediterranean Sea the species was also recorded off Cyprus (Katsanevakis et al., 2009), Turkey (Eryilmaz et al., 2003; Bilecenoglu, 2010) and Greece (Zachariou-Mamallinga & Corsini, 1994; Peristeraki et al., 2006).
The species is now considered as established in Greek waters (Zenetos et al., 2007) and waters off Malta (Schiberras & Schembri, 2006). After the first record of the blunthead pufferfish in the Ionian basin in 1991 (Tur-
si et al., 1992), this species showed a significant increase in abundance with time and there is now a steady population with the presence of spawning females (Ma-iorano et al., 2010).
According to Ragonese et al. (1992, 1997) the area between Sicily and Malta hosts an established population of the blunthead pufferfish. Ragonese et al. (1997) reported that 403 specimens of S. pachygaster were collected in the Sicily Strait in the period 1990-1994. Orsi Relini (2010) supposed that such a number of pufferfish probably arrived in the Sicily Strait before the date of the first record in the Mediterranean in the eighties (Oliver, 1981). In fact the area could probably be considered as a centre from which the population is spreading in different directions. However, nowadays, according to the data obtained by experimental surveys, the population in the waters of southern Sicily (and probably in the channel, too) is rather small (Ragonese & Murara, 2012).
Blunthead pufferfish in Adriatic waters
The first records of this species date from 1992 when Bello (1993) reported the occurrence of this species in southern Adriatic waters, in waters off Albania and close to Mola di Bari (Fig. 2). At about the same time Jardas & Pallaoro (1996) reported records of many blunthead pufferfish in 3 different localities in the southern part of the
Adriatic Sea: the Island of Susac (altogether 3 records with 5 specimens), Glavat (single record with 3 specimens) and Blitvenica (1 record with 1 specimen) (Jardas & Pallaoro, 1996). The previous northernmost extension of the range of the species was close to the very tip of the Istrian Penninsula - Cape Kamenjak (northern Adriatic Sea), when a specimen of blunthead pufferfish was captured on 8 November 1998 (Dulcic, 2002). The studied specimen from the waters off Piran represents the northernmost record of this species in the Adriatic and Mediterranean Sea, as well (Fig. 2).
In the Adriatic Sea three species of the family Tetrao-dontidae have been recorded to date (Tab. 2). In addition to the blunthead pufferfish, the other two species are Lagocephalus lagocephalus and L. sceleratus. The first is a Mediterranean species and it was first reported in the Adriatic Sea by Dulcic at Molunat in 2004 (Dulcic & Pallaoro, 2006). The second species is a Lessepsian migrant, which is rapidly spreading throughout the eastern Mediterranean and to other areas. The first record of this species is from November 2012, when a specimen was caught in waters off Dubrovnik (HINA, 2012).
Dispersal of S. pachygaster into the Mediterranean
There are two main hypotheses regarding the presence of the blunthead pufferfish in the Mediterranean Sea. The first deals with the recent immigration of the puffer-
Species	Locality	Country	Date	n	Depth (m)	TL (mm)	Source
Sphoeroides pachygaster	Sušac Island	Cro	16.3.1992	1			Jardas & Pallaoro, 1996
S. pachygaster	Sušac Island	Cro	11.4.1992	2		101, 120	Jardas & Pallaoro, 1996
S. pachygaster	Sušac Island	Cro	April 1992	2			Jardas & Pallaoro, 1996
S. pachygaster	Glavat Islet	Cro	August 1992	3	120	147	Jardas & Pallaoro, 1996
S. pachygaster	Blitvenica	Cro	15.11.1992	1	130-150	213	Jardas & Pallaoro, 1996
S. pachygaster	Mola di Bari	Ita	1992	some			Bello, 1993
S. pachygaster	S Adriatic Sea	Ita	1992	many	30-130		Bello, 1993
S. pachygaster	Albania	Alb	1992	1	85		Bello, 1993
S. pachygaster	Kamenjak (Pula)	Cro	8.11.1998	1	125	45	Dulcic, 2002
S. pachygaster	Budva	Mtg	5.1.2008	1	80	450	Joksimovic & Mandic, 2008
S. pachygaster	Šibenik	Cro	20.12.2008	1			Grubac, 2008
S. pachygaster	Piran	Slo	Nov 2012	1	20	348	This work
Lagocephalus lagocephalus	Molunat	Cro	2004	1	70	181	Dulcic & Pallaoro, 2006
L. sceleratus	Off Dubrovnik	Cro	Nov 2012	1		660	HINA, 2012
Tab. 2: Records of pufferfish species in the Adriatic Sea. Legend: Cro - Croatia, Ita - Italy, Alb - Albania, Mtg -Montenegro, Slo - Slovenia.
Tab. 2: Zapisi o pojavljanju različnih vrst rib napihovalk v Jadranskem morju. Legenda: Cro - Hrvaška, Ita - Italija, Alb - Albanija, Mtg - Črna gora, Slo - Slovenija
fish into the Mediterranean Sea from the Atlantic Ocean due to the gradual warming of the Mediterranean Sea. In fact the first records of this species originated only at the beginning of the eighties of the last century (Oliver, 1981). The second hypothesis is based on a painting by an Italian illustrator in 1558 in which the depicted fish could be identified as the blunthead pufferfish (Relini & Orsi Relini, 1995). The fish specimen, which was used as the model for the illustration came from the Delta of the Nile. Relini & Orsi Relini (1995) suggest that this fish was possibly present in the southern Mediterranean since ancient times. Obviously, since the opening of the Suez Canal was two centuries later, the blunthead pufferfish could not be related to Lessepsian migration.
Quero et al. (1998) studied the phenomenon of the occurrence of tropical fish species along the eastern Atlantic coast of Europe. According to their results there is a clearly visible trend of the spread of the blunthead pufferfish towards northern regions (from latitudes 40o to 55o) over the time period 1975-1995. These data clearly favour the first hypothesis of the recent immigration, related to water warming, into the Mediterranean Sea.
Many studies have confirmed that one of the main ecological parameters known to affect fish population is temperature (sensu Francour et al., 1994; Dulcic et al., 1999). Many thermophilous species of southern Mediterranean origin have been recently recorded in the Gulf of Trieste, such as Plectorhinchus mediterraneus (Lipej et al., 1996), Mola mola and Ranzania laevis, Luvarus imperialis and others (Lipej et al., 2007). Some alien fish species originating from the Indian Ocean were also reported, such as Siganus luridus (Poloniato et al., 2010) and Terapon theraps (Lipej et al., 2008).
The blunthead pufferfish is listed as a vulnerable species in the IUCN (Roberts, 1996); however its placement in this category probably needs a revision based on updated records. On the basis of available data we produced a map of the area with all records presented (Fig. 2). We tried to analyse the change in spatial pattern and its evolution over time in the Mediterranean Sea. For this purpose the reduced dataset were used, and year of first record within the 0.5 degree lat/long grid were analysed. For the whole study area the results indicate a statistically significant high clustering of the data and weak spa-
Fig. 3: Getis - Ord Gi Z scores indicating areas with significant spatial clustering of the first records in 0.5 degree lat/long greed. The red square represents the first record off the Atlantic coast of Europe.
Sl. 3: Getis - Ord Gi Z vrednost prikazuje statistično značilno gručanje prvih zapisov v 0,5-stopinjski mreži. Rdeči kvadrat označuje prvi zapis ob atlantski obali Evrope.
tial global autocorrelation (Morans I = 0.33, z = 3.61). Spatial autocorrelation occurs when the values of variables recorded at nearby locations are not independent from each other i.e. when adjacent geographic locations have very similar values (year of first records within the 0.5 degree grid in our case). Weak global spatial autocorrelation for the entire study area was expected and confirm the change in the spatial pattern of puffer fish records over time. It is common that the magnitude of spatial autocorrelation varies according to locations on a regional scale and exhibits significant clustering on a local scale. At local scale, Ord Gi statistics, with the search threshold for neighbours within 300 km, indicated few areas with statistically significant clustering. Figure 3 provides an overview of the calculated GiZ scores. As one can observe, high positive GiZ scores are clustered in the Adriatic, Aegean and Ionian Seas, indicating the directions of spread over the last two decades from the Sicily Channel (cluster with near zero values where the species had settled down and is constantly present over time). High negative values at the Atlantic coast of Europe and in the western Mediterranean represent the statistically significant cluster of earlier records and the direction of spread in the first decade after the very first record in 1978. It is quite evident that there is a clear eastward and northward spread of the blunthead pu-
fferfish. Similarly, temporal dynamics can be observed from Figure 2. The three directional distribution ellipses, calculated for three different time periods, also indicate the expected spatial trend in time. As it is clearly evident from Figures 2 and 3 the donor centre of spread has its origin in the Sicily Channel where the great majority of all findings of the blunthead puffer were recorded.
According to the available data many trends of the northward extension of the blunthead pufferfish can be seen. The rather evident trend of spread along the western coast of Europe and northward to Great Britain and Ireland has already been described by Quero et al. (1998).
In the Mediterranean Sea there are evident spread patterns in a north-easterly direction in the western Mediterranean, along the Levantine coast, in the Aegean Sea and finally in the Adriatic Sea, as well.
ACKNOWLEDGEMENTS
The authors wish to express their sincere thanks to Valter Ziza, Head of the Piran Aquarium, for providing us with a specimen of blunthead pufferfish. Special thanks go also to Vladimir Bernetic, who provided us with the available scientific literature on the matter. Special thanks also go to Prof. Thomas Mutz for his help and to two anonymous referees.
APPENDIX 1: Records of blunthead pufferfish in the eastern North Atlantic and the Mediterranean Sea Legend: Country: Por - Portugal, Esp - Spain, Ire - Ireland, Ita - Italy, GBr - Great Britain, Fra - France, Isr - Israel, Cro -Croatia, Mal - Malta, Gre - Greece, Alb - Albania, Tun - Tunisia, Alg - Algeria, Mtg - Montenegro, Slo - Slovenia, Cyp - Cyprus, Tur - Turkey. Region: ATL - Atlantic, W - West Mediterranean, e - east Mediterranean, c - central Mediterranean.
DODATEK 1: Pojavljanje ribe napihovalke v vzhodnem severnem Atlantiku in Sredozemskem morju. Legenda: Država - Por - Portugalska, Esp - Španija, Ire - Irska, Ita - Italija, GBr - Velika Britanija, Fra - Francija, Isr - Izrael, Cro - Hrvaška, Mal - Malta, Gre - Grčija, Alb - Albanija, Tun - Tunizija, Alg - Alžirija, Mtg - Črna gora, Slo - Slovenija, Cyp - Ciper, Tur - Turčija. Regija - ATL - Atlantik, W - zahodno Sredozemlje, e - vzhodno Sredozemlje, c - osrednje Sredozemlje
N	Locus	Country	Region	Date	n	Source
1	Estuary of the river Tago	Por	ATL	Jun1931	1	Gongalves, 1941
2	Nazaré'	Por	ATL	May 1978	1	Calvário et al., 1980
3	Sintra, 34 Nm NW of Cap Roca	Por	ATL	Jun1979	1	Calvário et al., 1980
4	Ribadesella	Esp	ATL	25.2.1980	1	Ortea et al., 1981
5	Cala Ratjada, Mallorca	Esp	W	1979	1	Oliver, 1981
6	Bay de Donegal	Ire	ATL	Jan 1984	1	Wheeler & van Oijen, 1985
7	Comarca del Garraf, Catalunya	Esp	w	1984	1	Cerro & Portas, 1984
8	Sicilian Channel	Ita	w	1985	2	Vacchi & Cau, 1985
9	Cala Ratjada, Mallorca	Esp	w	1984	1	Moreno & Roca, 1984
10	SW Spain Almeria	Esp	w	1986	2	Crespo et al., 1986
11	Gibraltar, Ceuta	Esp	w	1986	3	Crespo et al., 1986
12	Gulf of Cagliari, Sardinia	Ita	w	1986	5	Vacchi & Cau, 1986
13	San Remo	Ita	w	1986	1	Barletta & Torchio, 1986
14	Plymouth	GBr	ATL	1.2.1987	1	Quigley& Flannery, 1992
15	NW Sicilia	Ita	w	Jul 1988	1	Arculeo et al., 1994
16	Northern Spain	Esp	ATL	22.11.1988	1	Quéro et al., 1997
17	sud Gascogne	Esp	ATL	21.12.1988	1	Quéro et al., 1997
18	Lorient	Fra	ATL	2.5.1989	1	Quéro et al., 1997
19	La Rochelle	Fra	ATL	29.5.1989	1	Quéro et al., 1997
20	Concarneau	Fra	ATL	25.8.1989	1	Quéro et al., 1997
21	SW Ireland	Ire	ATL	19.8.1989	1	Quigley, 2002
22	Dingle Bay SW Ireland	Ire	ATL	18.10.1989	1	Quigley, 2002
23	Alboran Sea	Esp	w	12.6.1905	?	Camiñas et al., 1990
24	Imperia	Ita	w	1990	1	Fiorentino & Zamboni, 1990
25	coast of Israel	Isr	e	1990	1	Golani, 1996
26	Amendolara, Gulf of Taranto	Ita	c	1991	1	Tursi et al., 1992
27	Ashdod	Isr	e	1991	1	Psomadakis et al., 2006
28	Mola di Bari	Ita	a	1992	some	Bello, 1993
29	south Adriatic sea	Ita	a	1992	many	Bello, 1993
30	Gallipoli, Apulia	Ita	c	May 1992	1	Matarrese et al., 1996
31	Susac Island	Cro	a	11.4.1992	5	Jardas & Pallaoro, 1996
32	islet Glavat	Cro	a	22.8.1992	3	Jardas & Pallaoro, 1996
33	Blitvenica	Cro	a	15.11.1992	1	Jardas & Pallaoro, 1996
34	Ligurian sea	Ita	w	1992	?	Ragonese et a/., 1992
35	Malta	Mal	c	1992		Ragonese et a/., 1992
36	Lindos, Rodos	Gre	e	23.11.1992	1	Zachariou-Mamallinga & Corsini, 1994
37	Plimmiri, south of Lindos	Gre	e	24.11.1992		Zachariou-Mamallinga & Corsini, 1994
38	Lindos, Rodos	Gre	e	Mar 1993	10	Zachariou-Mamallinga & Corsini, 1994
39	Gulf of Gabes	Tun	c	1993	1	Bradai et a/., 1993
40	Albania	Alb	a	1993	1	Bello, 1993
41	Malta	Mal	c	1994		Schiberras & Schembri, 2006
42	Gulf de Lion	Fra	w	14.6.1995	1	Quignard & Raibaut, 1993
43	Strait of Sicily	Ita	w	1990-1994	403	Ragonese et a/., 1997
44	Elba	Ita	w	8.8.1996	1	Bedini, 1998
45	cape Kamenjak, Istra	Cro	a	8.11.1998	1	Dulcic, 2002
46	Saros bay TR	Tur	e	Oct 1999	1	Eryilmaz et a/., 2003
47	Serifos Island	Gre	e	2000	1	Zenetos et a/., 2007
48	South of Meganissi Island	Gre	e	2000	?	Zenetos et a/., 2007
49	Bozcaada Island	Tur	e	May 2001	1	Eryilmaz et a/., 2003
50	SW Ireland	Ire	ATL	1.1.2002	1	Quigley, 2002
51	SE Sikinos	Gre	e	26.8.2003	1	Peristeraki et a/., 2006
52	North of Lefkas Island	Gre	e	2004	?	Zenetos et a/., 2007
53	North of Lefkas Island	Gre	e	2005	?	Zenetos et a/., 2007
54	SE Serifos	Gre	e	14.6.2005	1	Peristeraki et a/., 2006
55	Cyprus	Cyp	e	2005	1	Katsanevakis et a/.,2009
56	SE Sikinos	Gre	e	12.6.2005		Peristeraki et a/., 2006
57	Elba	Ita	w	Aug 2005	?	Ligas et a/., 2006, 2007
58	SW Leros	Gre	e	4.7.2006	1	Peristeraki et a/., 2006
59	Serifos	Gre	e	Oct 2005		Zenetos et a/., 2007
60	SW Astypalaia	Gre	e	11.7.2006	1	Peristeraki et a/., 2006
61	Annaba	Alg	w	2008	1	Hemida et a/., 2009
62	off Budva	Mtg	a	5.1.2008	1	Joksimovic & Mandic, 2008
63	N of Corfu	Gre	e	16.3.2009	1	lianaskerkyra.blogspot. com/2009/03/strange-fish-in- corfu-waters.html
64	Tyrrhenian sea	Ita	w	2008-2009	1	Guerriero et a/., 2010
65	Gulf of Hammamet	Tun	c	2009		Chérif et a/., 2010
66	Karatasas coast, Iskenderun Bay	Tur	e	26.10.2010	1	Eleftheriou et a/., 2011
67	Samandag, Iskenderun Bay	Tur	e	28.10.2010	1	Eleftheriou et a/., 2011
68	Strait of Messina	Ita	w	Mar 2012	1	Giordano et a/., 2012
69	Piran	Slo	a	Nov 2012	1	This paper
NOVI NAJSEVERNEJŠI ZAPIS O POJAVLJANJU RIBE NAPIHOVALKE, SPHOEROIDES PACHYGASTER (OSTEICHTHYES: TETRAODONTIDAE) V SREDOZEMSKEM MORJU
Lovrenc LIPEJ & Borut MAVRIČ
Morska Biološka Postaja, Nacionalni Inštitut za biologijo, SI-6330 Piran, Fornače 41
E-mail: lipej@mbss.org
Dejan PALISKA
Univerza v Ljubljani, Fakulteta za pomorstvo in promet, SI-6320 Portorož, Pot pomorščakov 4
POVZETEK
Primerek ribe napihovalke vrste Sphoeroides pachygaster (Müller & Troschel, 1848) je bil ujet v vodah pred Piranom 22. novembra 2012. Gre za prvi zapis o pojavljanju te ribe v slovenskih vodah, obenem pa tudi za najsevernejši zapis o pojavljanju te vrste tako v Jadranskem kot tudi v Sredozemskem morju. Intenzivnejše pojavljanje te vrste v Sredozemskem morju se je začelo leta 1979 po prihodu iz Atlantskega oceana, trideset let kasneje pa se je napihoval-ka pojavila v najsevernejšemu predelu Sredozemskega morja. V prispevku avtorji na podlagi zbranih razpoložljivih podatkov o pojavljanju ribe napihovalke razpravljajo o širjenju areala te vrste v smeri proti vzhodu in severu.
Ključne besede: riba napihovalka, Sphoeroides pachygaster, širjenje proti severu, Jadransko morje
REFERENCES
Anselin, L. (1995): The local indicators of spatial association - LISA. Geogr. Anal., 27, 93-115.
Arculeo, M., S. Riggio & G. D'Anna (1994): First record of Sphoeroides pachygaster (Tetraodontidae) in the south Tyrrhenian (N/W Sicily). Cybium, 18 (2), 209-210.
Barletta, G. & M. Torchio (1986): Segnalazione di Bathypterois Günther e di Sphoeroides Lacèpede in ac-que imperiesi (Mar Ligure) (Osteichthyes). Quad. Civ. Staz. Idrobiol. Milano, 13, 31-34.
Bedini, R. (1998): First record of Sphoeroides pachygaster (Tetraodontidae) from the northern Tyrrhenian Sea. Cybium, 22 (1) 94-96.
Bello, G. (1993): Il pesce palla Sphoeroides cuta-neus nel mare Adriatico. Mem. Biol. Mar. Oceanogr., 18, 75-77.
Bilecenoglu, M. (2010): Alien fishes from Turkey -an updated review. In: Golani, D. & B. Appelbaum-Go-lani (eds.): Fish invasions of the Mediterranean Sea: Change and renewal. Pensoft Publishers, Sofia-Moscow, p. 189-217.
Bradai, M. N., M. Ghorbel & A. Bouain (1993): Premières observations dans le golfe de Gabès (Tunisie) de Sphoeroides cutaneus (Tétraodontidae). Cybium, 17 (1), 86.
Calvario, J. R., J. C. Marques & M. A. Pousada (1980): Occurrence of Sphoeroides pachygaster (Günther, 1870) (Pisces: Tetraodontidae) off the Portuguese coasts. Arquivos Mus. Bocage (2 ser.), 7 (10), 131-138.
Caminas, J. A., F. Ramos, J. C. Nunez & J. Baro (1990): Catalogue faunistique des especes capturees par
la flotille artisanale dans la mer d'Alboran (SE de l'Espagne). Rapp. Comm. int. Mer. Médit., 32 (1), p. 247.
Cerro, L. & F. Portas (1984): Addicio a la ictiofauna de la comarca del Garraf (Catalunya). Bull. Soc. Cat. Ic-tio. Herp., 9, 36-42.
Chérif, M., M. Ben Amor, M. Bdioui, S. Ben Salem, H. Missaoui & C. Capapé (2010): Additional records of the Blunthead puffer, Sphoeroides pachygaster (Osteichthyes: Tetraodontidae) off the Tunisian coast (Central Mediterranean). Annales, Ser. Hist. Nat., 20 (1), 1-6.
Crespo, J., J. G. Rey & A. Garcia (1986): New records of Sphoeroides pachygaster (Günther, 1870) (Pisces, Tetraodontidae) and Lophotes lacepede Giorna, 1809 (Pisces, Lophotidae) in the Mediterranean. Rapp. Comm. int. Mer Medit., 30 (2), p. 223.
Dragičevic, B. & J. Dulčic (2010): Fish invasions in the Adriatic Sea. In: Golani, D. & B. Appelbaum-Golani (eds.): Fish invasions of the Mediterranean Sea: Change and renewal. Pensoft Publishers, Sofia-Moscow, p. 255-266.
Dulčic, J. (2002): Northernmost occurrence of Sphoeroides pachygaster (Tetraodontidae) in the Adriatic Sea. Bull. Mar. Sci., 70 (1), 133-139.
Dulčic, J. & A. Pallaoro (2006): First record of the oceanic puffer (Lagocephalus lagocephalus lagocepha-lus Linnaeus, 1758), for the Adriatic Sea. J. Appl. Ichthyol., 1, 94-95.
Dulčic, J. & B. Dragičevic (2011): Nove ribe Jadranskog i Sredozemnog mora (New fishes of the Adriatic and the Mediterranean Sea). Institut za oceanografi-ju i ribarstvo, Državni zavod za zaštitu prirode, Split, 160 pp.
Dulcic, J., B. Grbec & L. Lipej (1999): Information on the Adriatic ichthyofauna - effect of water warming? Acta Adriat. 40 (2), 33-43.
Dulcic, J., A. Pallaoro & L. Lipej (2003): Lessepsian fish migrants reported in the Eastern Adriatic Sea. Annales, Ser. Hist. Nat., 13 (2), 137-144.
Eleftheriou, A., E. Anagnostopoulou-Visilia, E. Anastasopoulou, S. A. Ate$, N. El I. Bachari, L. Cavas,
C.	Cevik, M. Ulha, F. Cevik, A.-L. Delos, O. B. Derici, D. Erguden, N. Fragopoulu, A. Giangrande, T. Göksan, C. Gravili, M. Gurlek, A. Hattour, K. Kapiris, P. Kouraklis, S. Lamouti, E. Prato, L. Papa, G. Papantoniou, I. Parla-piano, D. Poursanidis, C. Turan & D. Yaglioglu (2011): New Mediterranean Biodiversity Records (December 2011). Medit. Mar. Sci., 12 (2), 491-508.
Eryilmaz, L., M. Özulug & N. Meri? (2003): The Smooth Pufferfish, Sphoeroides pachygaster (Tetraodon-tidae), new to the Northern Aegean Sea. Zoology in the Middle East, 28, 125-126.
Fiorentino, F. & A. Zamboni (1990): Ritrovamento di Epigonus constanciae (Giglioli, 1880) e nuove catture di Sphoeroides pachygaster (Günther, 1870) in Mar Ligure. Oebalia (Suppl.), XVI, 659-661.
Francour, P., C. F. Boudoresque, J. G. Harmelin, M. L. Harmelin-Vivien & J.-P. Quignard (1994): Are the Mediterranean waters becoming warmer? Mar. Pollut. Bull., 28 (9), 523-526.
Gabrosek, J. & N. Cressie (2002): The effect on attribute prediction of location uncertainty in spatial data. Geogr. Anal., 34, 262-285.
Getis, A. & J. K. Ord (1992): The Analysis of spatial association by use of distance statistics. Geogr. Anal., 24, 198-206.
Giordano, D., A. Profeta, L. Pirrera, F. Soraci, F. Per-dichizzi, S. Greco, A. Perdichizzi & P. Rinelli (2012): On the occurrence of the blunthead puffer, Sphoeroides pachygaster (Osteichthyes: Tetraodontidae), in the Strait of Messina (Central Mediterranean). Journal of Marine Biology, vol. 2012, article ID 462407. doi:10.1155/2012/462407
Golani, D. (1996): The marine ichthyofauna of the eastern Levant: history, inventory and characterization. Isr. J. Zool., 42, 15-55.
Golani, D., L. Orsi-Relini, L. Massutí & J. P. Quignard (2002): CIESM Atlas of Exotic species in the Mediterranean, Vol. 1. Fishes. Briand, F. (ed). CIESM Publishers, Monaco, 256 p.
Gon?alves, B. C. (1941): Colegao oceanográfica de
D.	Carlos I. Catálogo dos peixes. Trav. St. Biol. Marit. Lisb., 46, 108 pp.
Grubac, J. (2008): Otrovna napuhaca ulovljena blizu Sibenika. Slobodna Dalmacija, 21.12.2008.(http:// www.slobodnadalmacija.hr/Dubrovnik/tabid/75/article-Type/ArticleView/articleId/35254/Default.aspx)
Guerriero, G., A. Di Finizio & G. Ciarcia (2010): Biological pollution: molecular identification of non-native species in the Central Tyrrhenian Sea. Catrina, 5 (1), 41-47.
Hardy, G. S. (1981): New records of pufferfishes (family Tetraodontidae) from Australia and New Zealand, with notes on Sphoroides pachygaster (Müller & Troschel, 1848) and Lagocephalus sceleratus (Gmelin). Rec. Natl. Mus. N.Z., 1, 311-316.
Hemida, F., M. M. Ben Amor & C. Capape (2009): First confirmed record of the blunthead puffer, Sphoeroides pachygaster (Osteichthyes: Tetraodontidae) off the Algerian coast (southwestern Mediterranean). Pan-American Journal of Aquatic Sciences, 4 (2), 188-192.
HINA (2012): Nova vrsta ribe u Jadranskom moru. Novi list, Rijeka, 5.11.2012.
Hollowed, A. B. (1992): Spatial and temporal distribution of Pacific hake, Merluccius productus, larvae and estimate of survival during the early life stage. CalCOFI Report 33, p. 100-123.
Jardas, I. (1996): Jadranska ihtiofauna. Skolska knjiga. Zagreb, 533 pp. (In Croatian)
Jardas, I. & A. Pallaoro (1996): The record of Sphoeroides cutaneus (Günther, 1870) (Pisces, Tetraodontidae) in the Adriatic Sea. Oebalia, XXII, 83-90.
Joksimovic, A. & S. Mandic (2008): New fish species in the south Adriatic (Montenegrin coast) - Lessepsian migrant. Proceedings of 37th Annual Conference of the Yugoslav water pollution control society »Water 2008«. Mataruska banja, 36 June 2008, p. 297-300. (In Serbian) Katsanevakis, K., G. Tsiamis, G. loannou, N. Michai-lidis & A. Zenetos (2009): Inventory of alien marine species of Cyprus (2009). Medit. Mar. Sci., 10 (2), 109-133.
lianaskerkyra.blogspot.com/2009/03/strange-fish-in-corfu-waters.html
Ligas, A., R. Sirna & R. Sartor (2006): Prima segna-lazione di Sphoeroides pachygaster (Muller & Troschel, 1848) (Pisces, Tetraodontidae) nel Mar Tirreno Setten-trionale. Biol. Mar. Mediterr., 13, 274-275.
Ligas, A., P. Sartor, M. Sbrana, R. Sirna. & S. De Ra-nieri (2007): New Findings of Fistularia commersonii Ruppell, 1835 and Sphoeroides pachygaster (Muller & Troschel, 1848) in the northern Tyrrhenian Sea. Atti Soc. toscana Sci. nat. (Mem. B) 114, 131-133.
Lipej, L. & J. Dulcic (2004): The current status of Adriatic fish biodiversity. In: Griffiths, H. I., B. Krystufek & J. M. Reed (eds.): Balkan biodiversity: pattern and process in the European hotspot. Kluwer Academic, Dordrecht, London, p. 291-306.
Lipej, L., M. Spoto & J. Dulcic (1996): Plectorhin-chus mediterraneus from off north east Italy and Slovenia - the first records of the fish of the family Haemul-idae from the Adriatic Sea. J. Fish Biol., 48, 805-806.
Lipej, L., Z. Dobrajc, C. Castellarin, R. Odorico & J. Dulcic (2007): New records of some rare and less-known fishes in the Gulf of Trieste (Northern Adriatic). Annales, Ser. Hist. Nat., 17 (2), 171-176.
Lipej, L., B. Mavric, V. Ziza & J. Dulcic (2008): The largescaled terapon Terapon theraps: a new Indo-Pacific fish in the Mediterranean Sea. J. Fish Biol., 73, 18191822.
Maiorano, P., L. Sion, R. Carlucci, F. Capezzuto,
A.	Giove, G. Costantino, M. Panza, G. D'Onghia & A. Tursi (2010): The demersal faunal assemblage of the north-western Ionian Sea (central Mediterranean): current knowledge and perspectives. Chem. Ecol., 26 (Sup-pl.), 219-240.
Massutí, E., M. Valls & F. Ordines (2010): Changes in the western Mediterranean ichthyofauna: signs of tropicalisation and meridianizations. In: Golani, D. &
B.	Appelbaum-Golani (eds.): Fish invasions of the Mediterranean Sea: Change and renewal. Pensoft Publishers, Sofia-Moscow, p. 293-312.
Matarrese, A., G. D'Onghia, A. Tursi & M. Basani-si (1996): New information on the ichthyofauna of the South-Eastern Italian coasts (Ionian Sea). Cybium, 20 (2), 197-211.
Moreno, I. & I. Roca (1984): Second record of Spho-eroides cutaneus (Günther, 1870) (Tetraodontidae) from the Mediterranean Sea. Misc. Zool., 8, 301-303.
Oliver, P. (1981): Sobre la aparición de algunos pe-sces raros en las islas Baleares. Bol. Inst. Esp. Oceanogr., 6, 59-64.
Ord, J. K. & A. Getis, (1995): Local spatial autocorrelation statistics: distributional issues and an application. Geogr. Anal., 27, 286-306.
Orsi Relini, L. (2010): Non native marine fish in Italian waters. In: Golani, D. & B. Appelbaum-Golani (eds.): Fish invasions of the Mediterranean Sea: Change and renewal. Pensoft Publishers, Sofia-Moscow, p. 267-292.
Ortea, J., J. Alcazar, J. Carrasco & A Vizcaino (1981): Contribución al conocimiento de los peces marinos de Asturias. I. Notas sobre Liosaccus cutaneus (Günther), Echiodon dentatus Thompson y Centholophus niger (Gmelin) Boletín de Ciencias de la Naturaleza, 27, 163-173.
Peristeraki, P., G. Lazarakis, C. Skarvelis, M. Geor-giadas & G. Tserpes (2006): Additional records on the occurrence of alien fish species in the eastern Mediterranean Sea. Medit. Mar. Sci., 7 (2), 61-66.
Poloniato, D., S. Ciriaco, R. Odorico, J. Dulcic & L. Lipej (2010): First record of the dusky spinefoot Siganus luridus (Rüppell, 1828) in the Adriatic Sea. Annales, Ser. Hist. Nat., 20 (2), 161-166.
Psomadakis, P. N., P. Ceddia P. & M. Vacchi (2006): Additional record of Sphoeroides pachygaster (Pisces: Tetraodontidae) in the Tyrrhenian Sea and notes on the distribution of the species in the Mediterranean. JMBA2 - Biodiversity Records, 5186. (Published on-line)
Quéro, J.-C., M. H. Du Buit & J.-J. Vayne (1997): Les Captures De Poissons À Affinités Tropicales Le Long Des Côtes Atlantiques Européennes. Ann. Soc. Sci. Nat. Charente-Marit., 8 (6), 651-673.
Quéro, J.-C., M. H. Du Buit & J.-J. Vayne (1998): Les observations de poissons tropicaux et le réchauffement des eaux dans l'Atlantique européen. Oceanol. Acta, 21 (2), 345-351.
Quigley, D. T. G. & K. Flannery (1992): Further specimens of blunthead head puffer fish Sphoeroides
pachygaster Muller & Troschel 1848 (5. cutaneus Gun-ther1870) from Irish waters. Iri. Nat. J., 24, 155-157.
Quigley, D. (2002): Pufferfish in Irish waters. Sherkin Comment, 32, p. 4.
Quignard, J. P. & J. A. Tomasini (2000): Mediterranean fish biodiversity. Biol. Mar. Mediterr., 7 (3), 166.
Ragonese, S., G. Rivas & P. Jereb (1992): Spreading of the puffer 5phoeroides pachygaster Günther, 1870) (Pisces: Tetraodontidae) in the Sicilian Channel. Is it an »exploding« population? Rapp. Comm. int. Mer Médit., 33, p. 308.
Ragonese, S., P. Jereb & U. Morara (1997): Morpho-metric relationships of 5phoeroides pachygaster (Pisces: Tetraodontidae) of the Strait of Sicily (Mediterranean Sea). Cah. Biol. Mar., 38, 283-289.
Ragonese, S. & U. Morara (2012): Evidence of short-term discard mortality of trawled 5phoeroides pachygaster (Osteichthyes, Tetraodontidae) off the southern coast of Sicily (Central Mediterranean Sea). Pan-American Journal of Aquatic Sciences, 7 (2), 73-76.
Relini, M. & L. Orsi Relini (1995): Pesci palla in Mediterraneo, presenze antiche e recenti. Biol. Mar. Mediterr., 2 (2), 509-511.
Roberts, C. (1996): Liosaccus pachygaster. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. www.iucnredlist.org (21.10.2013)
Sciberras, M. & P. J. Schembri (2006): A critical review of records of alien marine species from the Maltese Islands and surrounding waters (Central Mediterranean). Medit. Mar. Sci., 8 (1), 41-66.
Tortonese, E. (1986): Tetraodontidae. In: Whitehead, P. J. P., M.-L. Bauchot, J.-C. Hureau, J. Nielsen & E. Tortonese (eds.): Fishes of the north-eastern Atlantic and the Mediterranean, Vol. 3. UNESCO, Paris, p. 1341-1347.
Tursi, A., G. D'Onghia, & A. Matarrese (1992): First finding of 5phoeroides pachygaster (Muller & Troschel, 1848) (Tetraodontidae) in the Ionian Sea (middle eastern Mediterranean). Cybium, 16 (2), 171-172.
Vacchi, M. & A. Cau (1986): The occurrence of 5phoeroides pachygaster (Günther, 1870) (Pisces, Tetraodontidae) in the Middle-West Mediterranean Sea. Cybium, 10 (2), 199-203.
Wheeler, A. & M. J. P. van Oijen (1985): The occurrence of 5phoeroides pachygaster (Osteichthyes- Tetra-odontiformes) off North-West Ireland. Zool. Meded., 59 (11), 101-107.
Zachariou-Mamalinga, H. & M. Corsini (1994): The occurrence of the fish 5phoeroides pachygaster in the south-eastern Aegean Sea (Greece). Ann. Mus. Goulan-dris, 9, 479-483.
Zenetos, A., V. Vassilopoulou, M. Salomidi & E. Pour-sanidis (2007): Additions to the marine alien fauna of Greek waters (2007 update). Marine Biodiversity Records 1, e91, doi.org/10.1017/S1755267207009281.