Section ECOLOGY AND ENVIRONMENTAL STUDIES

COMPARATIVE PLANKTON DYNAMICS IN ARABIAN

GULF AND SEA OF OMAN AT OPPOSITE SIDES OF THE
STRAIT OF HORMUZ

Prof. Waleed Hamza 1

Ms. Muzna Al Junaibi 2
Prof. Sergey Piontkovski 3
Dr. Khaled Al Hashmi 4
1, 2
Biology Department, College of Science, United Arab Emirates University
3, 4
Sultan Qaboos University, Sultanate of Oman

ABSTRACT
Phytoplankton and zooplankton monthly samples collected from the coastal
water stations, located in Ras Al Khaima Emirate (United Arab Emirates) and Sohar
governorate (Sultanate of Oman), during 2018-2019, have shown great variations,
not only in their community structures, but also in their species abundances.
Plankton samples were collected via vertical hauls from 6 m depth to the surface at
Ras Al Khaima, while in Sohar, samples were collected from 20 m depth to the
surface. The sample analyses revealed the dominance of diatoms during the warmer
months at both sample sites, with an increase of dinoflagellates during the colder
months, especially at Sohar. The abundances of certain species at each site, could
not be explained by the grazing of zooplankton (dominated by calanoid copepods)
on species-specific populations of phytoplankton, nor by seasonal temperature
variations. The hydrological regime at the Strait of Hormuz, separating the two
sample sites, as well as the differences in morphometric features and other
environmental parameters, could account for the ecological differentiation in
planktonic successions at both locations.
Keywords: Arabian Gulf, Sea of Oman, Planktonic succession, Hydrological
regime

INTRODUCTION
Algae bloom episodes recorded simultaneously in the Arabian Gulf (AG) and
the Sea of Oman (SO) during the last decade have led researchers to believe that
these two water bodies, which are connected by the Strait of Hormuz, have similar
environmental and ecological features. Detailed studies of planktonic community
structures and parallel surveys and analyses of phytoplankton and zooplankton
species-specific compositions on either side of the Strait of Hormuz are lacking
from literatures. Previous records of algae blooms in the AG and SO in 2008-2009
studied the blooming species and its intensity characteristics [1], [2] in the Arabian
Gulf region, and its expansion to the coastal areas at the Sea of Oman. The
icthyotoxic dinoflagellate species Cochlodinium polykrikoides was the cause of the
bloom and fish-kill phenomenon in the AG and SO, especially at Ras Al Khaima
(AG) and Fujirah (SO) emirates (both within the United Arab Emirates, UAE).
Previous algae blooms in the Sea of Oman by the dinoflagellate species Noctiluca
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miliaris were recorded by [3], [4], in the SO and at Kuwait Bay (AG). Recent
repeated Noctiluca scintillans blooms in the SO have been scantily recorded, and
equivalent blooms in the AG have in some cases not been recorded at all [5], [6],
and [7].
Water exchange between the AG and the SO through the Strait of Hormuz has
long been known. It is characterized by a deeper saline layer (39-40 ppt) outflowing
from the AG to the SO, and surface inflow of less saline (36-37 ppt) water from the
SO to the AG [8]. The inflow and outflow of water through the Strait of Hormuz is
unstable and does not follow the general circulation patterns of either the AG or the
SO [9], [8], and [10]. The fluctuation in the water trajectories between the two
basins is mainly controlled by wind direction and its duration may extend 2-8 days.
Moreover, the speed of the inflow surface current to the AG along the Iranian side
is 10 cm s-1, while the outflow deeper current to the SO is slower and reaches 3 m
s-1 [11]. This important feature, together with the coastal morphometry of the two
basins, impacts on the ecology, not only on the pelagic ecosystems, but also on sub-
pelagic and benthic systems. The average water depth in the AG is around 36 meters
and the maximum depth is about 100 meters at the Strait of Hormuz down to the
sandy bottom sediments. The average water depth in the SO is around 250 meters,
with maximum depth reaching 3000 meters down to the fine sandy and silty bottom
sediments [12].
General characteristics of phytoplankton and zooplankton communities in the
AG and SO are described in literature separately and for different periods, and are
summarized by [7]. During 2009-2011, diatoms contributed 70% to the total
phytoplankton abundance followed by dinoflagellates at 21% in the AG, while
small flagellates and diatoms contributed 10 and 25%, respectively, in the SO. For
another period [13] in the SO, the contribution of small flagellates and
dinoflagellates ranged between 25-40%. Zooplankton densities were found to be 10
times higher in the SO compared to the AG, however, the AG zooplankton
community was more diverse (210 species) compared to the 144 species identified
in the SO [14]. Copepods are the dominant species in both AG and SO zooplankton
communities, with seasonal peaks in winter and early summer seasons in the AG,
and multiple peaks in the SO, where there are monthly fluctuations in copepod
species abundances.
The present study was carried out to investigate the phytoplankton and
zooplankton dynamics at opposite locations in the AG and SO, with the aim of
identifying the relationships between planktonic community structures at these
locations, and taking account of the environmental and hydrological features
representing these marine basins.

MATERIALS AND METHODS

Studied stations, sampling and analytical procedures
The present study was carried out in locations in the Arabian Gulf (AG) and
the Sea of Oman (SO), separated by the Strait of Hormuz, during the period April
2018 through to May 2019. The selected station in the AG was in the coastal waters
of the Emirate of Ras Al Khaima (UAE), with geographic coordinates
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 Section ECOLOGY AND ENVIRONMENTAL STUDIES

25°47'22.31"N, 55°56'35.52"E. While, the selected station in the SO was in the

coastal waters of Sohar Governorate, at location 24°21'51.48"N, 56°44'48.52"E
(Figure 1).

Figure 1. Map of the sampled stations at the AG and SO ( ) indicating the

bathymetric features of the two basins (Modified from [17]).
During the study period, environmental parameters (water temperatures, water
salinities, dissolved oxygen and water pH) were measured in-situ within the water
column from which phytoplankton and zooplankton samples were collected.
Phytoplankton and zooplankton samples from Ras Al Khaima station (AG) were
collected at monthly intervals by vertical haul with a 20-µm nylon net from 6 m
depth to the surface. A similar procedure was followed for zooplankton sampling
but with an 80-µm nylon net. At the Sohar station (SO), due to water depth
differences and euphotic zone expansion, plankton samples were collected using
similar net meshes from 20 m depth to the surface.
Phytoplankton and zooplankton samples from both stations were analyzed for
their densities, abundances, and their species composition to the lowest taxonomic
ranks. The analyses were guided by local, regional and international taxonomic
references. Taxonomic ranking of phytoplankton was confirmed by collaboration
with M.G. Kholodny Institute of Botany, Ukraine, and for zooplankton with
assistance of the Institute of Biology of the Southern Seas, Russia. All counts and
numbers are referred to a volume unit of one cubic meter.
Remotely sensed data
Monthly satellite images of chlorophyll concentrations (mg m-3) at the surface
water of the studied area, and wind and current vectors (speed and direction) were
collected as shared information from the GIS Center at Sultan Qaboos University
(Sultanate of Oman). The obtained images represented monthly values during the

15
studied period (2018-2019) and were compared with long-term data analyses from
2002 until 2019.

RESULTS
Monthly variations in average water quality parameters showed marked
variations between Ras Al Khaima (RAK) and Sohar stations during the study
period. For instance, in RAK, water temperature averages ranged between 34.80°C
in July 2018 and 21.77 °C in March 2019. Moreover, water salinity ranged between
41.88 ppt in July 2018 and 36.7 in August 2018. Dissolved oxygen averages ranged
between 4.93 mg.l-1 in October 2018 and 6.69 mg.l-1 in March 2019 mg.l-1, and pH
averages ranged between 8.3 in September 2018 and 7.4 in April May 2019 (Table
1). On the other hand, at Sohar station water temperature averages ranged between
29.09°C in October 2018 and 23.40°C in February 2019. Water salinity averages
never exceed 36.82 ppt during the entire study period. Dissolved oxygen
concentrations averages ranged between 5.12 mg.l -1 in November 2018 and 10.84
mg.l-1 in May 2019, while water pH averages ranged between 8.00 in August 2018
and 8.60 in both March and April 2019 (Table 1).
Table 1. Monthly averages of in-situ measured water quality parameters at both
RAK (AG) and Sohar (SO) stations during the study period (2018-2019)

The analyses of phytoplankton species composition at RAK and Sohar stations

has revealed a higher diversity of RAK compared to Sohar, in that species diversity
exceeded 60 species in RAK station during November 2018, compared with 44
species at Sohar station during the same month (Figures 2). Although a few monthly
samplings were missed during the study periods at both sites, the diversity of
phytoplankton species for almost every month was higher at the RAK station than
the Sohar station.

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 Section ECOLOGY AND ENVIRONMENTAL STUDIES

Figure 2. Monthly phytoplankton species numbers and their diversities at RAK

(left) and Sohar (right) stations during the study period (2018-2019).
The monthly abundances of phytoplankton groups were dominated by
bacillariophyta (Bac.) species, which represented >90% of the phytoplankton
community structure at RAK, followed by dinoflagellates (Dino) that only
contributed 11-17 % of the community during the months of September and
October, 2018. Other groups such as cyanobacteria and chlorophyta never reached
more than 5 % except in June 2018, when they contributed about 60% of the
phytoplankton community density at RAK (Figure 3). At the Sohar station,
cyanobacteria and chlorophyta species dominated the phytoplankton community
structure, with values exceed 80 % during the months of August, 2018, as well as
March and April, 2019. In contrast, bacillariophyta species were dominant in both
June, 2018, and February, 2019, while dinoflagellate species contributed only
marginally during November, 2018 (Figure 3).

Figure 3. Monthly phytoplankton group abundances at RAK (left) and Sohar

(right) stations during the study period 2018-2019.
Zooplankton species diversity almost always showed higher diversity at Sohar
(SO) compared to RAK (AG) stations. The number of species identified at Sohar
was almost 3 times higher (around 100 species in November, 2018) than those
identified at RAK (around 32 species in January, 2019). Similar results were found
for the density levels, where the density of zooplankton per m -3 at Sohar was >10
times greater than those counted per m-3 at RAK (figure 4 and 5).

17
Figure 4. Monthly zooplankton species numbers and their diversities at RAK (left)
and Sohar (right) stations during the study period (2018-2019).
In both stations, zooplankton identified groups belonged to the calanoids,
cyclopoids and harpactocoids. Calanoid species always dominated the Sohar
samples, while in RAK both calanoids and cyclopoids represented almost 80 % of
the monthly community structure (figure 4).
The relationship between phytoplankton biomass and zooplankton densities per
cubic meter at both stations was plotted by month (figure 5). Apart from the
differences in their biomasses and their densities, there was no match in seasonal
succession trends between the planktonic communities in the two sampling
locations. At the RAK station, the zooplankton community peaked in June, 2018,
followed by a sharp decline in September, 2018. After this, moderate densities
followed by another decline in March, 2019, and recovery in May, 2019, preceded
another peak in early summer. In March, 2019, the phytoplankton community
showed a marked peak in its biomass that followed a less marked one in December,
2018 (figure 5).

Figure 5. Monthly succession of phytoplankton biomass (mg.m -3) and zooplankton

density (ind.m-3*103) at RAK (left) and Sohar (right) stations during the study
period (2018-2019).
The zooplankton community at the Sohar station showed very high density
during July, 2018, that exceeded 4.5 million individuals per cubic meter. This high
density declined in the following months to reach its minimum in October, 2018,
with only six thousands (6,000) individuals per cubic meter. From November, 2018,
until April, 2019, there was a relatively limited increase in the densities during
December, 2018, and April, 2019, to reach only 600 thousand and 300 thousand
individuals, respectively. Surprisingly, the phytoplankton reached its high peak

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 Section ECOLOGY AND ENVIRONMENTAL STUDIES

biomass (2500 mg.m-3) during July, 2018, when zooplankton biomass also peaked
(figure 5). The other phytoplankton increased in biomass, but with much less
pronounced peak observed in January, 2019, to a value of around 800 mg.m -3.
The dominant species of phytoplankton and zooplankton showed no
complementary trends between the two basins, except for the individual case of the
bacillariophyta species Guinardia flaccida that was dominant in the Sohar
phytoplankton community during June, and dominated the RAK phytoplankton
community during the period from January to May, 2019. The Calanoid species
Temora turbinata that dominated the RAK zooplankton community only during
April, 2018, dominated the Sohar zooplankton community during August, 2018,
February, 2019 and April, 2019 (table 2).
Table 2. Monthly dominant species of both phytoplankton and zooplankton
communities identified in RAK (AG) and Sohar (SO) stations during the study
period (2018-2019).

The integration of wind components over the studies region during the period
May, 2003, to February, 2018 (NOAA/NCDC blend daily 0.25º- 10 m altitude),
especially during the months where similar planktonic organisms dominated their
communities structure, showed direction changes that may have affected surface
water movement directions and velocities. During other months, wind components
were analyzed separately for each basin. According to figure 6, wind direction was
from the AG towards the SO, through the Strait of Hormuz, during April, while in

19
August, the opposite direction prevailed. However, in January, a northwestern wind
may have prevent the movement of waters between the two basins.

Figure 6. Integrated wind directions over the studied region during the period
2003-2018 for the months of January (left), April (middle) and August (right).
Data Courtesy of NOAA-NCEL (modified from [10]).
The calculated wind stress data on the sea surface during 2019, at different
months, corresponded to the long-term integrated wind directions (Figure 7),
showing fluctuations in surface water interconnection between the two basins.

Figure 7. Calculated monthly sea surface anomalies over the studied region
during January (left), April (middle) and August (right) 2019.
Although satellite images of Chlorophyll a concentrations obtained during
2019 did not assay every month, the obtained results for the months of January and
April, 2019, correlate with the calculated wind data and sea surface anomalies
during the same months. Very low and dispersed chlorophyll a concentrations along
the Iranian coast were detected during January, 2019, while in April, 2019, high
concentrations of chlorophyll a in the coastal area of the Sea of Oman were
observed.

DISCUSSION
Both the Arabian Gulf and the Sea of Oman are known for their high
phytoplankton production, and their ample nutrient salts concentrations. That has
been regularly confirmed by historical data and in recent studies [11], [15], [16],
[17], [7], and [10]. However, the differences observed in the present study in the
phytoplankton community structures of these water bodies, could be mainly due to
the differences in their environmental parameters, namely, high water salinity and
high dissolved silicate contributed by dust storms over the AG have favored the
domination of the bacillariophyta group over the other phytoplankton [17]. On the
other hand, the Oceanic water salinity characterizing the SO may have allowed the
growth of other phytoplankton groups. Although dust was also deposited over the
SO, the superior area and depth of the SO basin may have decrease the availability
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 Section ECOLOGY AND ENVIRONMENTAL STUDIES

of dissolved silicates to bacillariophyta species, allowing other phytoplankton

groups to compete with them.
The summarized comparison between dominant phytoplankton groups in the
AG and the SO by [7], is confirmed in the present study, though with slight
differences in percentages between the two community structures of the studied
stations. Moreover, the high densities of zooplankton communities observed in this
study in the Sohar station have exceeded by a factor of 15 the densities at the RAK
station, which surpassed by a factor of 10 the results reported by [14], when the
zooplankton densities at both basins are compared. Although slight differences in
temperatures and the possible adaptation of plankton to such variations may be
factors, the distinct dynamics of planktonic seasonal successions between the
studied stations are best explained by atmospheric and hydrologic regime
differences between the studied areas. It has been reported that the SO surface water
is dominated by oceanic water that flows along the Iranian coast and mixes with
cool advected water during the winter northeast monsoon, while during the summer
southwest monsoon, an upwelling persists along the south coast of Oman [10]. [10],
also noted that conditions off the northern coast of Oman are mainly controlled by
the advected water, the strong heating during summer, the impact of an intense eddy
field, the outflow of the Arabian Gulf, and variable wind directions. Furthermore,
[7] mentioned that, the AG water outflow spreads in the form of a subsurface
salinity maximum (occupying a depth range between 200 and 350 meters). They
also found that seasonal variations of atmospheric temperatures create atmospheric
pressure anomalies, which could be one of the factors mediating the water mass
transport between the AG and SO. These findings, representing a general pattern,
may confirm the contribution of wind stresses over the sea surface and water current
flow directions between the two basins. In our study, the plankton samples were
collected from 6 meters depth from RAK station and from 20 meters depth at Sohar
station. These represent the upper mixed layers in the two basins, when compared
to their average depths. At such layer passive transportation of planktonic
organisms by water currents and the presence of atmospheric anomalies during
different months (figures 6 and 7), may result in dispersal of planktonic
communities and may also disrupt and/or dilute their structures. By linking the
water exchange features and the differences in the bathymetry and the hydrology of
the two basins [9], and [7], irregularities in seasonal trends of planktonic
successions are expected. Furthermore, absence of harmonization in seasonal
productivity peaks, as well as species dominance similarities, may result from
surface water movement anomalies that lead to random but regular and harmonized
periods where algae blooms occur simultaneously in the two basins.
In the present study, the dominance of the bacillariophyta species Guinardia
flaccida at Sohar station in June, 2018, and its dominance in the RAK
phytoplankton community from January, 2019, until May, 2019 (table 2), could be
a result of the different hydrological processes associated with the entry of SO
surface water into the AG through the Strait of Hormuz. This surface water flow
follows the anticyclonic gyre front in Qatari waters [9] and [8]. The return flow of
waters through the RAK station may carry the same species. On the other hand, the
dominance of the calanoid copepod Temora turbinate of the RAK zooplankton

21
community during April, 2018, and its dominance in Sohar during August, 2018, as
well as in February and April, 2019 could also be an indication of possible sharing
of dominant species between the two basins, though at different seasons or different
years. This could be the result of water exchange between the two basins and
sporadic dispersal of certain planktonic species, due to their high densities and their
passive transportation during water circulation.
By observing the relationship between phytoplankton and zooplankton
communities in the present study at RAK station, it is evident that there is an
ecological succession that characterizes its planktonic community dynamics. The
high peaks of the zooplankton density period are always accompanied by low
phytoplankton biomass and vice versa (figure 5). In contrast, at the Sohar station,
equivalent correlation between phytoplankton and zooplankton communities is
quite absent. Indeed, the zooplankton density peak observed in July, 2018, was
accompanied by a peak in phytoplankton biomass (Figure 5). These features, lead
us to conclude that the RAK (AG) zooplankton community is able to control the
phytoplankton biomass, while at Sohar (SO), both zooplankton and phytoplankton
communities are governed by other factors. In their study, [18], [7] and [10]
mentioned that the SO is characterized by high fish landings, with major
contribution of the filter feeder sardine and juvenile fish that may play substantial
role in controlling planktonic communities. By this means, the planktonic
interactions at Sohar station would be governed mainly by predation-prey
interactions, while at RAK, grazing interactions are the key feature of its ecological
succession. This conclusion provides an improved model to explain the effects of
hydrological interactions between the AG and SO stations that are separated by the
Strait of Hormuz. It also expands our understanding of simultaneous algae bloom
phenomena in the two basins.

CONCLUSION
This study was carried out during 2018-2019 in the form of detailed monthly
investigations of planktonic communities and relevant environmental parameters at
two stations: the Arabian Gulf (RAK) and the Sea of Oman (Sohar), on opposite
sides of the Strait of Hormuz. The study shows that both morphometric and
hydrologic differences between the two basins play major roles in controlling their
planktonic community structures and their seasonal dynamics. It is concluded that
the ecological succession in the Arabian Gulf is characterized by grazing of
zooplankton on phytoplankton, while predation-prey interaction more appropriately
describes the ecological succession in the Sea of Oman.

ACKNOWLEDGEMENTS
This research funded by the SQU-UAEU Collaborative Research Grant
CL/SQU-UAEU/ /18/04 and 31S321). We would like to thank Y. Bryantseva (M.G.
Kholodny Institute of Botany, Ukraine) and E. Popova (Institute of Biology of the
Southern Seas, Russia) for their assistance in identifying and confirming the
taxonomy of phytoplankton and zooplankton samples respectively. Our gratitude
extended to Dr. S. Al-Ghais and Dr. S. Varadharajulu from EPDA Ras Al Khaima

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 Section ECOLOGY AND ENVIRONMENTAL STUDIES

Government, for their approval and assistance in sampling and samples processing
at RAK station and for Mr. S. Al-Khusaibi, H. Al-Habsi. B. Al-Bawaiqi and H. Al-
Lawati for their technical assistance in sampling of Sohar station and in data
processing for the Sea of Oman. A special thanks to Dr. A. Fowler for his careful
revision of the manuscript editing.

REFERENCES
[1] Richlen M.L., Morton S.L., Jamali E.A., Rajan A., and Anderson D.M., The
catastrophic 2008-2009 red tide in the Arabian Gulf region, with observations on
the identification and phylogeny of the fish-killing dinoflagellate Cochlodimium
polykrikoides. Harmful Algae, Vol. 9, pp 163-172, 2010.
[2] Zhao, J., Ghedira, H., Monitoring red tide with satellite imagery and
numerical models: a case study in the Arabian Gulf. Mar. Poll. Bull. Vol.79, pp
305-313, 2014.
[3] Glibert P., Eutrophication and harmful blooms: a complex global issue,
examples from the Arabian Sea including Kuwait Bay, and an introduction to the
global ecology and oceanography of harmful algal blooms (GEOHAP) program.
International Journal of Oceans and Oceanography, Vol. 2, pp 157-169, 2014.
[4] Gomes, H. R.; Goes J.I. Matondkar S.G. Parab S.G. Al-Azri A. and Thoppil
P.G., Blooms of Noctiluca miliaris in the Arabian Sea-An in situ and satellite study.
Deep Sea Research, Part I, Vol. 55, pp 751 765, 2008.
[5] Zhang S., H. Liu, C. Guo and Harrison P., Differential feeding and growth
of Noctiluca scintillans on monospecific and mixed diets. Marine Ecology Progress
Series, Vol. 549, pp 27-40, 2016.
[6] Harrison, P.J., Piontkovski, S.A., Al-Hashmi, K.A., Overview of decadal
ecosystem changes in the Western Arabian Sea and the occurrence of algal blooms.
Journal of Agricultural and Marine Sciences. Vol. 23, pp 11-23, 2019.
[7] Piontkovski S.A., Hamza W., Al-Abri N., Al-Busaidi S., and Al-Hashmi,
K.A., A comparison of seasonal variability of Arabian Gulf and the Sea of Oman
pelagic ecosystems. Aquatic Ecosystem Health and Management, Vol. 22, pp 108-
130, 2019.
[8] Wang, Z., DiMarco, S.F., Jochens, A.E., and Ingle S., High salinity in the
northern Arabian Sea and Sea of Oman. Deep-Sea Research (I) Vol. 74, pp 14-24,
2013.
[9] Pous, S.P., Carton, X., and Lazure, P., Hydrology and circulation in the
Strait of Hormuz and the Gulf of Oman- Results from the GOGP99 Experiment: 1.
Strait of Hormuz. Journal of Geophysical Research, 109 Vol. 12 - Sect. 3; pp.
C12037. 2004.
[10] Al-Hashmi, K.A., Piontkovski, S.A., Bruss, G., Hamza, W., Al-Junaibi,
M., Bryantseva Y., and Popova, E., Seasonal variations of planktonic communities
in coastal waters of Oman. Int. J. Oceans and Oceanography. Vol. 13, pp 395-426,
2019.
23
 [11] Reynolds R.M., Physical oceanography of the Gulf, Strait of Hormuz, and
the Gulf of Oman results from the Mt Mitchell expedition. Marine Pollution
Bulletin, Vol. 27, pp 35-59, 1993.
[12] Hamza, W., Munawar, M., Protecting and managing the Arabian Gulf:
past, present and future. Aquat. Ecosyst. Health Manag. Vol. 12, pp 429-439, 2009.
[13] Polikarpov I., Saburova M., Al-Yamani F., Diversity and distribution of
winter phytoplankton in the Arabian Gulf and the Sea of Oman. Continental Shelf
Research, Vol. 19, pp 85-99, 2016.
[14] ROPME, State of the marine environment report. ROPME/GC-16/1-ii.
Regional Organization for the Protection of the Marine Environment. Kuwait, pp 1-
225, 2013.
[15] Shriadah, M.A., Nutrient salts in the United Arab Emirates waters (the
Arabian Gulf and the Gulf of Oman). Pakistan J. Mar. Sci. Vol. 15, pp 1-9, 2006.
[16] Emara, H.I., Nutrient salts, inorganic and organic carbon contents in the
waters of the Persian Gulf and the Gulf of Oman. J. Pers. Gulf. Vol. 1, pp 33-44,
2010.
[17] Hamza, W., Enan, M.R., Al-Hassini, H., Stuut, J-B., de-Beer, D., Dust
storms over the Arabian Gulf: a possible indicator of climate changes consequences.
Aquat. Ecosyst. Health Manag. Vol. 14, pp 260-268, 2011.
[18] Piontkovski S.A., Al-Oufi H.S., Al-Jufaili S., Seasonal and interannual
changes of sardine landings in the Sea of Oman. Marine Fisheries Review, Vol. 76-
3, pp 50-59, 2014.

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