INTRODUCTION
Undeterred by Forbes’ (1844: 170) notorious
observation “Zero of Animal Life probably about 300
fathoms”, the scientific exploration of the Levantine
bathyal biota commenced with the extensive voyages
of the Pola (1890-1893). In 1910 the Danish
Oceanographic Expedition to the Mediterranean,
aboard the Thor, sampled nine sites along the western
limits of the Levantine Sea, from Cyrenaica to
THE BATHYAL FAUNA OF THE LEVANTINE SEA
63
SCI. MAR., 68 (Suppl. 3): 63-72
S
CIENTIA
M
ARINA
2004
MEDITERRANEAN DEEP-SEA BIOLOGY. F. SARDÀ, G. D’ONGHIA, C.-Y. POLITOU and A. TSELEPIDES (eds.)
The limit of the sea: the bathyal fauna of
the Levantine Sea*
BELLA S. GALIL
National Institute of Oceanography, Israel Oceanographic and Limnological Research, P.O.B. 8030,
Haifa 31080, Israel. E-mail: bella@ocean.org.il
SUMMARY: In the present study, the fish, molluscs, crustaceans and echinoderms collected at depths between 734 and 1558
m during a series of cruises conducted between 1988 and 1999 off the coast of Israel, supplemented by a photographic sur-
vey carried out southwest of Cyprus at a depth of 2900 m, were analysed. The main objectives were to determine the faunal
composition of the bathybenthic assemblages in the southeastern Levantine Sea, and to compare them with the western
Mediterranean assemblages in order to elucidate whether general trends in their bathymetric distribution and population den-
sity may be related to environmental/geographic factors. Considering the sampling effort, the diverse gear used and the
extended period of sampling, we may assume that the low number of species and specimens recorded actually reflects a low-
diversity, low-density deep water fauna. The faunal scarcity may cause a different parcelling of the populations which is
reflected in bathymetric distributions that in many cases extend to greater depths than in the Western Mediterranean. The
Levantine bathybenthos is composed of autochthonous, self-sustaining populations of opportunistic, eurybathic species that
have settled there following the last sapropelic event.
Key words: Mediterranean Sea, Levantine Sea, bathyal, diversity, abundance, bathymetric distribution, new records, fish,
Decapoda, Amphipoda, Cumacea, Mollusca.
RESUMEN: E
L LÍMITE DEL MAR
:
LA FAUNA BATIAL DEL MAR
L
EVANTINO
. – En este estudio se analizan los peces, los molus-
cos, los crustáceos y los equinodermos recolectados a profundidades entre 734 y 1558 m, durante una serie de campañas de
muestreo realizadas entre 1988 y 1999 delante de las costas de Israel. Estas muestras se complementan con fotografías sub-
marinas realizadas a 2900 m de profundidad al suroeste de Chipre. El objetivo es determinar la composición faunística de
las asociaciones batipelágicas en el sureste del mar Levantino y compararla con las asociaciones del Mediterráneo occiden-
tal, con el fin de observar si existen tendencias generales en su distribución batimétrica y en las densidades de las pobla-
ciones, de manera que puedan ser relacionadas con factores geográficos y ambientales. Considerando el esfuerzo realizado
en muestreo, los diversos artes usados y el amplio período de muestreo, podemos asumir que el bajo número de especies y
especímenes registrado refleja realmente la baja diversidad y la baja densidad de la fauna de las aguas profundas. La escasa
fauna puede causar un distinto parcelado de las poblaciones que se refleja en su distribución batimétrica la qual en muchos
casos se extiende hacia las más grandes profundidades del Mediterráneo occidental. El batibentos levantino esta compuesto
por especies autóctonas, poblaciones de especies oportunistas auto-sostenibles y euribáticas, que colonizaron estos fondos
desde el último evento sapropélico.
Palabras clave: mar Mediterráneo, mar Levantino, batial, diversidad, abundancia, distribución batimétrica, nuevas citas,
peces, Decapoda, Amphipoda, Cumacea, Mollusca.
*Received June 9, 2003. Accepted January 27, 2004.
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Rhodes. However, as
that expedition was concerned
mainly with plankton investigations, and “dredgings
were only occasionally made … the result will thus
necessarily be but poor” (Stephensen, 1915). The
Lamont Geological Observatory research vessel Vema
collected few deep benthic samples in 1958 (Barnard,
1964). Four deep-water shark species were collected
in five hauls using bottom long-line at depths of 1330-
1440 m off the Mediterranean coast of Israel in 1985
(Golani, 1987). The German research vessel Meteor
[cruise 5] in 1987, collected benthic samples at sites
between Crete and Israel at depths ranging between 95
and 4396 m during a ten-day sojourn (Janssen, 1989).
The handful of deep sea fishes gathered persuaded
Klausewitz (1989) that “this collecting trip confirmed
the paucity of the deep sea fish fauna in the eastern
basin of the Mediterranean”. The Meteor returned in
1993 [cruise 25], to collect deep-sea biota from the
region between Crete, Cyprus and Egypt at depths
ranging between 194 and 2812 m, as well as sea bed
litter (Galil et al., 1995). The fauna of Eratosthenes
Seamount, south of Cyprus, was briefly sampled by
the Poseidon [cruise 201/2] in February 1994 (Galil
and Zibrowius, 1998). Among the faunistic explo-
rations of the Levantine bathyal, only Gilat and
Gelman (1984), and Jones in Priede and Bagley
(2000) made use of photographic equipment to
observe the deep fauna in its environment.
The meagre number and extent of the explo-
rations prompted Fredj and Laubier (1985), in their
review on the deep Mediterranean benthos, to
declare that “...the southern part of the Levant Sea
has practically never been studied”.
The present study analysed the data obtained
from samples collected by a box corer and a bottom
trawl fitted with a plankton net during a series of
cruises conducted between 1988 and 1999 as part of
pollution monitoring surveys by the Israel
Oceanographic and Limnological Research (IOLR)
off the coast of Israel, at depths between 734 and
1558 m, supplemented by a photographic survey
carried out southwest of Cyprus, at a depth of 2900
m. The main objectives were to determine the faunal
composition of the bathybenthic assemblages in the
southeastern Levantine Sea, and to compare them
with the western Mediterranean assemblages in
order to elucidate whether general trends in their
bathymetric distribution and population density may
be related to environmental factors. Although
restricted to a small area and to the depth range men-
tioned above, this study has significantly increased
our knowledge of the Levantine deep sea fauna.
STUDY AREA AND METHODS
The Levant Sea occupies the easternmost
Mediterranean, east of the line connecting Rhodes
and the coast of Cyrenaica. It is isolated from the
deep Atlantic waters by the topographical and
hydrological barriers posed by the shallow Gibraltar
Straits and the Siculo-Tunisian sill. The Levantine
deep water mass is distinguished by salinity and
temperature values that are higher than in the rest of
the Mediterranean (T < 13.8ºC, S < 38.74, below
700 m) (Hecht et al., 1988). The Levantine Sea is
ultra-oligotrophic (Berman et al., 1984; Krom et al.,
1991): Chlorophyll a concentrations are as low as
0.4 µg l
–1
nearshore, and decrease offshore to 0.05
µg l
–1
(Berman et al., 1986; Yacobi et al., 1995).
The area investigated is located off the coast of
Israel, at depths between 734 and 1558 m. The mate-
rial was collected during monitoring surveys of two
deepwater waste-dumping sites: an acidic sludge dis-
posal site off Haifa (between 33°00’N 34°37’E and
33°01’N 34°47’E), a coal fly ash disposal site off
Hadera (between 32°38’N 34°02’E and 32°36’N
34°16’E), and a control site off Atlit (between
32°53’N 34°10’E and 32°51’N 34°23’E). The sam-
ples were collected aboard the R/V Shikmona (720
HP; 27 m), using a modified Agassiz benthic trawl
(2.3 m width and 0.9 m height), a 45 ft Marinovich-
type deep water trawl (codend mesh 6 mm) with a 0.5
mm plankton net secured atop, and a 0.062 m
2
box-
corer with an effective penetration of 40 cm (Ocean
Instruments model 700 AL) (Table 1). The samples
were preserved in 10% buffered formalin aboard ship.
In the laboratory, the samples were washed and
sieved through a 500 µm mesh, preserved in 70%
alcohol, stained in Rose Bengal and sorted.
The photographic survey took place southwest of
Cyprus, at depth of 2900 m. A total of 80 hours of
video were recorded utilising a remote operating
vehicle (Remora 6000, Phoenix).
RESULTS
Fishes
A total of 566 specimens identified to 31 species
were collected during the study. Four species were
new records for the eastern Mediterranean, four
species were new records for the Levantine Sea and
five species were newly recorded from the Israeli
coast (Galil and Goren, 1994; Goren and Galil,
64
B.S. GALIL
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