Background

The red pandora, Pagellus bellottii Steindachner, 1882, is a tropical and subtropical demersal Sparidae (Porgies) that lives over hard or sandy bottoms in coastal marine waters. It mostly occurs at depths ranging from 10 to 50 metres. It is distributed in the eastern Atlantic, from the Strait of Gibraltar to Angola and the Canary Islands where it is occasionally recorded (Bauchot and Hureau, 1986; Franqueville, 1983). The species is also recorded in the southwestern Mediterranean, in the Alboran Sea, off the Algerian coasts and in the Gulf of Gabes (Oral, 2010). The records of the species in the eastern Mediterranean, Syrian and Israeli waters (Fricke, et al. 2014, Fig. 1) is questioned. We consider these records are misidentification of Pagrus pagrus (Linnaeus, 1758). Pagellus bellottii was also included in the checklist of the fishes from Portugal as a consequence of records coming from the Algarve region, about 36°59′ N, 8° W, the northernmost records in Atlantic waters (de Castro, 1967; Erzini et al., 1996; Carneiro et al., 2014; Carneiro com. pers.). Some specimens were also recorded in Spanish Mediterranean waters, in the Bay of Almería about 36°47′ N, 2°25′ W (Lucena, et al. 1982), the northernmost records for the Mediterranean waters. Herein, the first record of P. bellottii in the Bay of Biscay is reported.

Results

SYSTEMATICS

Order PERCIFORMES

Family SPARIDAE Rafinesque, 1810

Genus Pagellus Valenciennes, 1830

Pagellus bellottii Steindachner, 1882

(Figure 1)

Fig. 1
figure 1

Pagellus bellottii Steindachner, 1882, voucher MNHN-IC 2015–0144, 265 mm TL, 26 Oct. 2014, southern Bay of Biscay, France. Scale bar = 10 cm

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The two specimens were similar in size, with a total length (TL) of 265 and 274 mm respectively (Fig. 1). Morphometrics, weights and meristic counts are presented in Table 1. Diagnostic characters of the two specimens included occipital scales extending forward to the anterior margin of eye; molar teeth in two series; anal fin with 10 rays; bright silvery pale red colours; dark red mark at the origin of the lateral line and the upper margin of opercle; fins yellow-pinkish; inside of mouth whitish; and longitudinal yellow and grey stripes along body.

Table 1 Morphometrics (mm), weight (g) and meristic counts of the two specimens of Pagellus bellottii from the Bay of Biscay
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The average seasonal water temperature cycle at the depth of 20 m in the southeast Bay of Biscay in 2005–2012 was overall cooler than those of the corresponding seasons to the south of Angola in 1955–1964, although the annual cycles of the two areas overlap (Fig. 2). Recent winter temperatures in the Bay of Biscay have been cooler by more than 2 °C than winter temperatures to the south of Angola in 1955–1964, while summer temperatures are comparable. Winter temperatures in the Bay of Biscay are almost 5 °C cooler than in the Gulf of Cádiz.

Fig. 2
figure 2

Comparison of the average seasonal temperature cycle in the South eastern Bay of Biscay in 2005–2012 (thick black line) to that from 1955–1964 in the Gulf of Cádiz (blue line) and to the South of Angola by 16–17 °S (thin continued line) and by 17–18 °S (dotted line). Sea temperatures from the World Ocean Atlas 2013 (Boyer et al. 2013)

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Discussion

The two current records are the northernmost occurrences ever reported for the species, and the first record in the Bay of Biscay and French waters. These new records represent a significant range extension for the species of about 6°45′ to the north. The specimens were collected together with congeners of the species Pagellus erythrinus (Linnaeus, 1758). Four species of Pagellus are now known to be present in French waters: P. acarne (Risso, 1827), P. bogaraveo (Brünnich, 1768), P. erythrinus and the newly recorded P. bellottii.

Several sparids have displayed a significant northward range extension in European Atlantic marine waters over the last few decades. The phenomenon was observed for Diplodus cervinus (Lowe, 1838), D. vulgaris (Geoffroy Saint-Hilaire, 1817), D. sargus (Linnaeus, 1758), and Pagrus auriga Cuvier, 1816 (Quéro et al., 2004; Bañón et al., 2014; pers. obs.). The population abundance of uncommon native sparid species in the Bay of Biscay, such as Sparus aurata Linnaeus, 1758, Dentex dentex (Linnaeus, 1758), and Boops boops Linnaeus 1758 (Quéro et al., 2009; pers. obs.), appears to have increased over the last few decades. The observed northward range extension of a number of slope and shelf species in the Bay of Biscay was shown to coincide with the warming of both the Atlantic waters of the poleward Slope Current, and that of shelf waters in the southern Bay of Biscay (Quéro et al., 1998). The warming of the Bay of Biscay over 40 years, from the mid-1960s to the mid-2000s, was shown to be roughly twice as fast as that of the overall North Atlantic basin (Michel et al., 2009), which could be seen as a factor in favour of the extension of a mostly tropical species such as P. bellottii. Nevertheless, even in the warmer southeast Bay of Biscay temperatures remain far cooler than in the tropical coastal waters of West Africa where the species is most abundant. In summer and only in surface layers, recent water temperatures in the Bay of Biscay are similar to or warmer than temperatures in the 1950s-1960s in the Strait of Gibraltar and South Angola, corresponding to extreme latitudes of the native range. In contrast, winter temperatures in the southeast Bay of Biscay are much cooler than in all other areas of the species native range and recent occurrence.

The status of the two individuals at hand is unknown. They may be either vagrant members of their population, as was recently observed for a cod Gadus morhua individual fished in the Mediterranean Sea (Morey et al., 2012), members of a small recently established resident population or seasonal migrants. In any case, whether larvae were carried or juveniles/adults actively migrated to the catching location is unknown. It seems unlikely that these two individuals were spawned in the Gulf of Cádiz, from which they would have had to have been carried/travelled more than 1300 km. The recent occurrences in the Gulf of Cádiz of both juveniles from 6 cm TL and mature individuals up to 30 cm TL may reflect the existence of a breeding population in this area (Torres et al., 2012; Carneiro et al., 2014). Furthermore, previous occurrences may have passed unnoticed in commercial catches and even in fishery science surveys, because of the strong similarity to other Pagellus species, and in particular to P. erythrinus. It is worth noting that it is only because two individuals of each species were caught in the same haul in 2014, that this first record was made. If individuals of P. bellottii were previously caught without P. erythrinus in the same haul, they may have been misidentified as the latter. Therefore, the confirmation of the status of the species from Portugal to the Bay of Biscay requires careful monitoring of Pagellus spp. catches in all sampling opportunities from the Gulf of Cádiz to the Bay of Biscay in forthcoming years.

Conclusions

The northern extension of the distribution range of the tropical-subtropical Sparidae Pagellus bellottii most probably is a consequence of the warming waters of the Bay of Biscay over the past decades.

Methods

On the 26th October 2014, two specimens of Pagellus bellottii were collected by the French research vessel Thalassa during the fishery science survey Evhoe 2014 at the station S-0853 (Fig. 3). They were collected by bottom trawl in the southeast of the Bay of Biscay, near the French coast, off Vieux-Boucau-les-Bains at 43°46.9′-43°48.9′ N, 1°27.2′-1°26.6′ W at 25.1–25.6 m depth. Species identification was made according to Bauchot and Hureau (1986). Fresh specimens were photographed in a standard manner then a tissue sample was taken from the specimens and recorded under collection numbers (BPS-3069 and 3070) for future genetic analyses. The specimens were fixed in 4 % formalin for some weeks then preserved in 70 % ethanol and deposited in the Ichthyologic Collection of the Muséum national d’Histoire naturelle of Paris (MNHN-IC 2015–0144 and 2015–0145).

Fig. 3
figure 3

Location (black dot) of the catch of two specimens of Pagellus bellottii off Vieux-Boucau-les-Bains, southern Bay of Biscay, France, on 26 Oct. 2014

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