A single adult female specimen of Halosaurus ovenii Johnson, 1864 was captured by trammel nets at a depth of about 200 m off the coast of Arbatax (Sardinia, Italy) in early April 2007. Macroscopic and microscopic analysis of the gonad showed a postspawning ovary. This is the fourth documented capture of this fish in the Mediterranean Sea, representing the north-easternmost record for this species in this geographic area. Furthermore, the present specimen was fished at the shallowest depth ever recorded before

Follesa, Maria Cristina

Merella, Paolo

Motomura, Hiroyuki

Pais, Antonio

Acta Ichthyologica et Piscatoria

English

Pais,A.

Merella,P.

Follesa,M.C.

Motomura,H.

ARPHA Preprints

North-easternmost record of Halosaurus ovenii (Actinopterygii: Notacanthiformes: Halosauridae) in the Mediterranean Sea, with notes on its biology

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UnissResearch

According to Froese and Pauly (2008), 10 species belong
to the genus Halosaurus Johnson, 1864 (Notacanthiformes:
Halosauridae), 3 of which commonly occur in the eastern
Atlantic: H. guentheri Goode et Bean, 1896; H. johnsoni-
anusVaillant, 1888; andH. ovenii Johnson, 1864. The lat-
ter species is a benthopelagic fish that lives at bathyal
depths ranging from 440 to 1700 m (Sulak 1986). It is an
amphiatlantic fish commonly present off Madeira, the
Canaries, the Azores and the western African coast from
Morocco to South Africa (Sulak 1990), but it was also
recorded off the coast of Portugal (Costa and
Reiner 1978). In the western Atlantic its geographical dis-
tribution is well documented from the Caribbean and the
Gulf of Mexico, and off the Lesser Antilles (Sulak 1990,
Smith 2002). This fish is extremely rare in the
Mediterranean Sea, where Golani et al. (2002) considered
it as an exotic species. Its first record in this basin was
reported by Dieuzeide (1963) from off the Habibas Islands
(Algeria, western Mediterranean) where, in 1960, a single
specimen was captured at 550 m depth. Cau and Deiana
(1979) documented the occurrence of this species off Capo
Teulada (Sardinia, Italy, central western Mediterranean) at
620 m depth. Finally, D’Onghia et al. (2004) captured one
specimen (229 mm TL) off the Balearic Islands, at 2800 m
depth, the deepest record for this species. The capture of
another specimen of H. ovenii off the eastern coast of
Sardinia represented a noteworthy opportunity to collect
additional information on its geographical distribution
and biology.
On 3 April 2007, one adult specimen of H. ovenii was
captured by trammel nets 1.5 miles off the port of Arbatax
(Sardinia, central western Mediterranean, lat 39°57.6'N,
long 09°44.3'E) on a muddy bottom at a depth of about
200 m (Fig. 1). The fish was photographed (Fig. 2), frozen
at –20°C, and in the laboratory it was identified to species
according to Sulak (1986).
Its stomach contents were weighed, examined, and then
stored in 70% ethanol. After sex determination (female),
the maturity stage of the ovary was assessed macroscopi-
cally according to Holden and Raitt (1974), and then the
gonad was fixed in 5% buffered formalin for subsequent
histological analysis. Afterwards, the ovary was dehydrat-
ed in an ascending ethanol series, embedded in resin, and
semi-thin sections of 3 mm were stained with sodium
iodoeosine and toluidine blue following Dominici’s
method modified for methacrylate (Mazzi 1977). The iden-
ACTA ICHTHYOLOGICA ET PISCATORIA (2009) 39 (1): 33–37 DOI: 10.3750/AIP2009.39.1.06
* Correspondence: Dr Antonio Pais, Sezione di Acquacoltura e Gestione delle Risorse Acquatiche, Dipartimento di Scienze Zootecniche, Università di Sassari,
Via E. De Nicola 9, 07100 Sassari (Italy), phone: +39 079 229345, fax: +39 079 229302, e–mail: pais@uniss.it
NORTH-EASTERNMOST RECORD OF HALOSAURUS OVENII (ACTINOPTERYGII:
NOTACANTHIFORMES: HALOSAURIDAE) IN THE MEDITERRANEAN SEA,
WITH NOTES ON ITS BIOLOGY
Antonio PAIS1*, Paolo MERELLA2, Maria Cristina FOLLESA3, and Hiroyuki MOTOMURA4
1 Sezione di Acquacoltura e Gestione delle Risorse Acquatiche, Dipartimento di Scienze Zootecniche,
Università di Sassari, Sassari, Italy
2 Sezione di Parassitologia e Malattie Parassitarie, Dipartimento di Biologia Animale,
Università di Sassari, Sassari, Italy
3Dipartimento di Biologia Animale ed Ecologia, Università di Cagliari, Viale Poetto 1, 09126 Cagliari, Italy
4 The Kagoshima University Museum, 1–21–30 Korimoto, Kagoshima 890–0065, Japan
Pais A., Merella P., Follesa M.C., Motomura H. 2009. North-easternmost record of Halosaurus ovenii
(Actinopterygii: Notacanthiformes: Halosauridae) in the Mediterranean Sea, with notes on its biology.
Acta Ichthyol. Piscat. 39 (1): 33–37.
Abstract. A single adult female specimen of Halosaurus ovenii Johnson, 1864 was captured by trammel nets at
a depth of about 200 m off the coast of Arbatax (Sardinia, Italy) in early April 2007. Macroscopic and micro-
scopic analysis of the gonad showed a postspawning ovary. This is the fourth documented capture of this fish in
the Mediterranean Sea, representing the north-easternmost record for this species in this geographic area.
Furthermore, the present specimen was fished at the shallowest depth ever recorded before.
Keywords: Halosaurus ovenii, new record, Mediterranean, Sardinia
tification of the oocyte developmental stages was carried
out using an optical microscope, according to the scale
proposed by Forberg (1982). Gonadic maturity stages
were determined on the basis of the relative abundance of
oocytes cells in the more advanced stage of development.
The diameter of nucleated oocytes was measured by
means of a graduated ocular.
Body surface, mouth, gills, body cavity and all inter-
nal organs were examined for metazoan parasites. Fresh
scrapings of the organs were also observed microscopical-
ly for protozoan parasites. Finally, the fish was deposited
in the ichthyological collection of the Department of
Animal Science at the University of Sassari (registration
code DSZ-04/07).
The fish was a female of 470 mm total length and 92.9 g
total weight. Morphometric and meristic characters of the
present specimen and of those formerly reported from the
Mediterranean are given in Table 1. The ovary appeared
flaccid and whitish in colour, with many visible eggs. It
was located just after the liver, composed by 2 distinct
lobes originating proximally at about 1/5 of its length and
extending posteriorly up to 3/4 of the body cavity.
Macroscopic analysis revealed that it was at a post-spawn-
ing stage (stage 7 of Holden and Raitt scale). Subsequent
histological analysis showed the presence of few mature
oocytes [translucent (T), mean diameter 940 ± 39.74 µm,
ripe egg stage by Forberg (1982)]; atresic oocytes (ATR);
and post-ovulatory follicles (POF) (Fig. 3). The stomach
and intestine contained only remains of organic matter and
neither protozoan nor metazoan parasites were detected.
Halosaurus ovenii was first captured in the
Mediterranean Sea only in 1960 (Dieuzeide 1963). This
Pais et al.34
Fig. 1.Map of records of Halosaurus ovenii in the Mediterranean. Black square = presently described record; hollow
squares = previous records
Fig. 2. Halosaurus ovenii, DSZ-04/07, 470 mm TL, Sardinia, Western Mediterranean, 200 m depth
late record probably led Quignard and Tomasini (2000) to
consider it as a recent migrant and not just a rare species
in this geographical area. In any case, the present capture
confirms its presence also in the central-western part of
the basin and corresponds to its north-easternmost occur-
rence in the Mediterranean. As the present specimen was
fished at the shallowest depth ever recorded before (i.e.,
about 200 m), this new record shows an extremely wide
bathymetric range for this species, i.e., from 200 m down
to 2800 m (D’Onghia et al. 2004). The female examined
appeared at a post-spawning stage, in accordance with the
observations reported by Cau and Deiana (1979) for
another H. ovenii specimen captured in the same period of
the year. The presence of POFs in the gonadic tissue was
a further confirmation that the ovary was quiescent
(Cinquetti and Rinaldi 1987, Yoneda et al. 1998).
The specimen examined was totally free of parasites.
There are few reports of parasites from members of the
family Halosauridae, the only ones are from Aldrovandia
oleosa Sulak, 1977 (cf. Sulak 1977) and Halosauropsis
macrochir (Günther, 1878) (cf. Klimpel et al. 2001 and
references therein, Klimpel et al. 2008), and no parasite
had been reported from Halosaurus spp. Moreover,
Boxshall (1998), in a broad study on host specificity
Halosaurus ovenii in the Mediterranean Sea 35
Fig. 3. Histological stage of Halosaurus ovenii ovary: translucent oocyte (T), post-ovulatory follicle (POF) and atresic
oocyte (ATR); scale bar = 250 µm
Character This study Dieuzeide(1963)
Cau and Deiana
(1979)
M
or
ph
om
et
ric
[m
m
]
Total length [TL] 470 — 520
Gnathoproctal length 21 — —
Body height 35 1/14 TL 38
Head length [HL] 56 1/8 TL       62.5
Snout length 23 — —
Predorsal length 142 — 157
Preanal length 225 — —
Eye diameter length [ED] 9.2–9.5 1/5 HL      13.3
Eye diameter height 6.5–6.7 —        8.0
Interorbital width 4.7 1/2 ED —
M
er
is
tic
Dorsal fin rays I+10 I+10 I+10
Anal fin rays >190 191 152
Pectoral fin rays I+14 I+10 I+12
Ventral fin rays I+7–8 I+9 I+9
Gill rakers 1st arch 12 (9+3) — —
Lateral line scales anterior to anus 66–67 — —
Pyloric caeca 11 12 11
Table 1
Morphometric and meristic data of Halosaurus ovenii specimens recorded in the Mediterranean Sea
in copepod parasites of deep-sea fishes, observed that
copepods had failed to colonise Halosaurus spp. from the
North Atlantic, in spite of their relative abundance in deep
sea habitat. The absence of parasites in the examined host
(although limited to a single specimen) seems to confirm
some drawback in the success of parasite on this genus,
notwithstanding these fish inhabit the bathyal grounds,
where the diversity of parasites is relatively high
(Marcogliese 2002, Klimpel et al. 2006). Finally, a further
mechanism that could explain the absence of parasites is
the enemy release hypothesis (ERH). It points out that
introduced species usually invade new areas without their
natural parasites or they lose them in the new habitat
(Colautti et al. 2004). This will suggest that H. ovenii
should be considered as exotic to the Mediterranean Sea
and/or that the examined host was caught at a shallower
depth of the natural range of the species. But caution
should be considered in the uncritical acceptance of ERH,
as recently reported by Pais et al. (2007, 2008) and
Merella et al. (2008), for Atlantic and Lessepsian
migrants from the same geographical area.
ACKNOWLEDGEMENTS
The authors are especially grateful to Silverio Calisi,
skipper of the F/V SBRIGLIO of Arbatax, to Mario and
Simone Demelas for providing the specimen and useful
information on the catch site, and to two anonymous review-
ers for their critical comments and valued suggestions.
REFERENCES
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Received: 17 May 2008
Accepted: 4 December 2008
Published electronically: 20 May 2009
Halosaurus ovenii in the Mediterranean Sea 37


North-easternmost record of <i>Halosaurus ovenii</i> Actinopterygii: Notacanthiformes: Halosauridae) in the Mediterranean Sea, with notes on its biology

A single adult female specimen of Halosaurus ovenii Johnson, 1864 was captured by trammel nets at a depth of about 200 in off the coast of Arbatax (Sardinia, Italy) in early April 2007. Macroscopic and microscopic analysis of the gonad showed a postspawning ovary. This is the fourth documented capture of this fish in the Mediterranean Sea, representing the north-easternmost record for this species in this geographic area. Furthermore, the present specimen was fished at the shallowest depth ever recorded before

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According to Froese and Pauly (2008), 10 species belongto the genus Halosaurus Johnson, 1864 (Notacanthiformes:Halosauridae), 3 of which commonly occur in the easternAtlantic: H. guentheri Goode et Bean, 1896; H. johnsoni-anusVaillant, 1888; andH. ovenii Johnson, 1864. The lat-ter species is a benthopelagic fish that lives at bathyaldepths ranging from 440 to 1700 m (Sulak 1986). It is anamphiatlantic fish commonly present off Madeira, theCanaries, the Azores and the western African coast fromMorocco to South Africa (Sulak 1990), but it was alsorecorded off the coast of Portugal (Costa andReiner 1978). In the western Atlantic its geographical dis-tribution is well documented from the Caribbean and theGulf of Mexico, and off the Lesser Antilles (Sulak 1990,Smith 2002). This fish is extremely rare in theMediterranean Sea, where Golani et al. (2002) consideredit as an exotic species. Its first record in this basin wasreported by Dieuzeide (1963) from off the Habibas Islands(Algeria, western Mediterranean) where, in 1960, a singlespecimen was captured at 550 m depth. Cau and Deiana(1979) documented the occurrence of this species off CapoTeulada (Sardinia, Italy, central western Mediterranean) at620 m depth. Finally, D’Onghia et al. (2004) captured onespecimen (229 mm TL) off the Balearic Islands, at 2800 mdepth, the deepest record for this species. The capture ofanother specimen of H. ovenii off the eastern coast ofSardinia represented a noteworthy opportunity to collectadditional information on its geographical distributionand biology.On 3 April 2007, one adult specimen of H. ovenii wascaptured by trammel nets 1.5 miles off the port of Arbatax(Sardinia, central western Mediterranean, lat 39°57.6'N,long 09°44.3'E) on a muddy bottom at a depth of about200 m (Fig. 1). The fish was photographed (Fig. 2), frozenat –20°C, and in the laboratory it was identified to speciesaccording to Sulak (1986).Its stomach contents were weighed, examined, and thenstored in 70% ethanol. After sex determination (female),the maturity stage of the ovary was assessed macroscopi-cally according to Holden and Raitt (1974), and then thegonad was fixed in 5% buffered formalin for subsequenthistological analysis. Afterwards, the ovary was dehydrat-ed in an ascending ethanol series, embedded in resin, andsemi-thin sections of 3 mm were stained with sodiumiodoeosine and toluidine blue following Dominici’smethod modified for methacrylate (Mazzi 1977). The iden-ACTA ICHTHYOLOGICA ET PISCATORIA (2009) 39 (1): 33–37 DOI: 10.3750/AIP2009.39.1.06* Correspondence: Dr Antonio Pais, Sezione di Acquacoltura e Gestione delle Risorse Acquatiche, Dipartimento di Scienze Zootecniche, Università di Sassari,Via E. De Nicola 9, 07100 Sassari (Italy), phone: +39 079 229345, fax: +39 079 229302, e–mail: pais@uniss.itNORTH-EASTERNMOST RECORD OF HALOSAURUS OVENII (ACTINOPTERYGII:NOTACANTHIFORMES: HALOSAURIDAE) IN THE MEDITERRANEAN SEA,WITH NOTES ON ITS BIOLOGYAntonio PAIS1*, Paolo MERELLA2, Maria Cristina FOLLESA3, and Hiroyuki MOTOMURA41 Sezione di Acquacoltura e Gestione delle Risorse Acquatiche, Dipartimento di Scienze Zootecniche,Università di Sassari, Sassari, Italy2 Sezione di Parassitologia e Malattie Parassitarie, Dipartimento di Biologia Animale,Università di Sassari, Sassari, Italy3Dipartimento di Biologia Animale ed Ecologia, Università di Cagliari, Viale Poetto 1, 09126 Cagliari, Italy4 The Kagoshima University Museum, 1–21–30 Korimoto, Kagoshima 890–0065, JapanPais A., Merella P., Follesa M.C., Motomura H. 2009. North-easternmost record of Halosaurus ovenii(Actinopterygii: Notacanthiformes: Halosauridae) in the Mediterranean Sea, with notes on its biology.Acta Ichthyol. Piscat. 39 (1): 33–37.Abstract. A single adult female specimen of Halosaurus ovenii Johnson, 1864 was captured by trammel nets ata depth of about 200 m off the coast of Arbatax (Sardinia, Italy) in early April 2007. Macroscopic and micro-scopic analysis of the gonad showed a postspawning ovary. This is the fourth documented capture of this fish inthe Mediterranean Sea, representing the north-easternmost record for this species in this geographic area.Furthermore, the present specimen was fished at the shallowest depth ever recorded before.Keywords: Halosaurus ovenii, new record, Mediterranean, Sardiniatification of the oocyte developmental stages was carriedout using an optical microscope, according to the scaleproposed by Forberg (1982). Gonadic maturity stageswere determined on the basis of the relative abundance ofoocytes cells in the more advanced stage of development.The diameter of nucleated oocytes was measured bymeans of a graduated ocular.Body surface, mouth, gills, body cavity and all inter-nal organs were examined for metazoan parasites. Freshscrapings of the organs were also observed microscopical-ly for protozoan parasites. Finally, the fish was depositedin the ichthyological collection of the Department ofAnimal Science at the University of Sassari (registrationcode DSZ-04/07).The fish was a female of 470 mm total length and 92.9 gtotal weight. Morphometric and meristic characters of thepresent specimen and of those formerly reported from theMediterranean are given in Table 1. The ovary appearedflaccid and whitish in colour, with many visible eggs. Itwas located just after the liver, composed by 2 distinctlobes originating proximally at about 1/5 of its length andextending posteriorly up to 3/4 of the body cavity.Macroscopic analysis revealed that it was at a post-spawn-ing stage (stage 7 of Holden and Raitt scale). Subsequenthistological analysis showed the presence of few matureoocytes [translucent (T), mean diameter 940 ± 39.74 µm,ripe egg stage by Forberg (1982)]; atresic oocytes (ATR);and post-ovulatory follicles (POF) (Fig. 3). The stomachand intestine contained only remains of organic matter andneither protozoan nor metazoan parasites were detected.Halosaurus ovenii was first captured in theMediterranean Sea only in 1960 (Dieuzeide 1963). ThisPais et al.34Fig. 1.Map of records of Halosaurus ovenii in the Mediterranean. Black square = presently described record; hollowsquares = previous recordsFig. 2. Halosaurus ovenii, DSZ-04/07, 470 mm TL, Sardinia, Western Mediterranean, 200 m depthlate record probably led Quignard and Tomasini (2000) toconsider it as a recent migrant and not just a rare speciesin this geographical area. In any case, the present captureconfirms its presence also in the central-western part ofthe basin and corresponds to its north-easternmost occur-rence in the Mediterranean. As the present specimen wasfished at the shallowest depth ever recorded before (i.e.,about 200 m), this new record shows an extremely widebathymetric range for this species, i.e., from 200 m downto 2800 m (D’Onghia et al. 2004). The female examinedappeared at a post-spawning stage, in accordance with theobservations reported by Cau and Deiana (1979) foranother H. ovenii specimen captured in the same period ofthe year. The presence of POFs in the gonadic tissue wasa further confirmation that the ovary was quiescent(Cinquetti and Rinaldi 1987, Yoneda et al. 1998).The specimen examined was totally free of parasites.There are few reports of parasites from members of thefamily Halosauridae, the only ones are from Aldrovandiaoleosa Sulak, 1977 (cf. Sulak 1977) and Halosauropsismacrochir (Günther, 1878) (cf. Klimpel et al. 2001 andreferences therein, Klimpel et al. 2008), and no parasitehad been reported from Halosaurus spp. Moreover,Boxshall (1998), in a broad study on host specificityHalosaurus ovenii in the Mediterranean Sea 35Fig. 3. Histological stage of Halosaurus ovenii ovary: translucent oocyte (T), post-ovulatory follicle (POF) and atresicoocyte (ATR); scale bar = 250 µmCharacter This study Dieuzeide(1963)Cau and Deiana(1979)Morphometric[mm]Total length [TL] 470 — 520Gnathoproctal length 21 — —Body height 35 1/14 TL 38Head length [HL] 56 1/8 TL       62.5Snout length 23 — —Predorsal length 142 — 157Preanal length 225 — —Eye diameter length [ED] 9.2–9.5 1/5 HL      13.3Eye diameter height 6.5–6.7 —        8.0Interorbital width 4.7 1/2 ED —MeristicDorsal fin rays I+10 I+10 I+10Anal fin rays >190 191 152Pectoral fin rays I+14 I+10 I+12Ventral fin rays I+7–8 I+9 I+9Gill rakers 1st arch 12 (9+3) — —Lateral line scales anterior to anus 66–67 — —Pyloric caeca 11 12 11Table 1Morphometric and meristic data of Halosaurus ovenii specimens recorded in the Mediterranean Seain copepod parasites of deep-sea fishes, observed thatcopepods had failed to colonise Halosaurus spp. from theNorth Atlantic, in spite of their relative abundance in deepsea habitat. The absence of parasites in the examined host(although limited to a single specimen) seems to confirmsome drawback in the success of parasite on this genus,notwithstanding these fish inhabit the bathyal grounds,where the diversity of parasites is relatively high(Marcogliese 2002, Klimpel et al. 2006). Finally, a furthermechanism that could explain the absence of parasites isthe enemy release hypothesis (ERH). It points out thatintroduced species usually invade new areas without theirnatural parasites or they lose them in the new habitat(Colautti et al. 2004). This will suggest that H. oveniishould be considered as exotic to the Mediterranean Seaand/or that the examined host was caught at a shallowerdepth of the natural range of the species. But cautionshould be considered in the uncritical acceptance of ERH,as recently reported by Pais et al. (2007, 2008) andMerella et al. (2008), for Atlantic and Lessepsianmigrants from the same geographical area.ACKNOWLEDGEMENTSThe authors are especially grateful to Silverio Calisi,skipper of the F/V SBRIGLIO of Arbatax, to Mario andSimone Demelas for providing the specimen and usefulinformation on the catch site, and to two anonymous review-ers for their critical comments and valued suggestions.REFERENCESBoxshall G.A. 1998. Host specificity in copepod parasites ofdeep-sea fishes. Journal of Marine Systems 15: 215–223.DOI: 10.1016/S0924-7963(97)00058-4.Cau A., Deiana A.M. 1979. Prima segnalazione di Halosaurusovenii Johnson, 1863 nei mari italiani. Quaderni della CivicaStazione Idrobiologica di Milano 7: 127–130. [In Italian.]Cinquetti R., Rinaldi L. 1987. Change in the gonadal histologyof Padogobius martensi (Pisces: Gobiidae) during the repro-ductive cycle. Italian Journal of Zoology 54: 233–241. DOI:10.1080/11250008709355589.Colautti R.I., Ricciardi A., Grigorovich I.A., MacIsaac H.J.2004. 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Check-list of the Fishes of the Eastern Tropical Atlantic (CLOFE-TA), Vol. 1. JNICT, Lisbon, SEI, Paris, UNESCO, Paris.Yoneda M., Tokimura M., Fujita H., Takeshita N.,Takeshita K., Matsuyama M., Matsuura S. 1998.Reproductive cycle and sexual maturity of the anglerfishLophiomus setigerus in the East China Sea with a note onspecialized spermatogenesis. Journal of Fish Biology 53:164–178. DOI: 10.1111/j.1095-8649.1998.tb00118.x.Received: 17 May 2008Accepted: 4 December 2008Published electronically: 20 May 2009Halosaurus ovenii in the Mediterranean Sea 37

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North-easternmost record of <i>Halosaurus ovenii</i> Actinopterygii: Notacanthiformes: Halosauridae) in the Mediterranean Sea, with notes on its biology

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