Nature and Science 2010;8(5)

ENVIRONMENTAL STUDIES ON THE MUDSKIPPERS IN THE

INTERTIDAL ZONE OF KUWAIT BAY
Bahija E. Al-Behbehani* & Hussain M. A. Ebrahim
*
Science Department, College of Basic Education, PAAET, Kuwait
2-College of Health Sciences, PAAET, Kuwait
bshm7000@[Link]

Abstract: This work deals with monitoring mudskippers in their natural environment (intertidal zone) along the
Kuwait Bay muddy shores in the State of Kuwait. This is to provide information is concerning the environmental
factors effecting mudskipper diversity in Kuwait Bay. Kuwait Bay is a large mud-flat with a fascinating associated
fauna of mud-skippers and crabs provide rewarding feeding-grounds for many birds. A number of fifty mudskipper
samples are collected during the hot summer season (July and August) of the year 2009 and examined for parasites
and to evaluate the different environmental factors controlling the biodiversity in this marine environment. The
results of the present study indicate the abundance of the mudskippers allover the intertidal mud flat of the Bay and
the total absence of either external and/or internal parasites in the mudskipper tissues and organs. Mudskippers are
found to be completely amphibious fish that are adapted to live in the intertidal environment. Mudskippers are very
active when they are outside the water, feeding and interacting with one another. The mud in the Kuwait Bay
environment is very good for burrowing in, since the particles are very sticky, unlike sand. Often, the mudskipper
form mixed colonies with digging crabs (Fiddler crabs-Caidae). Specific physiological and behavioural changes in
bioindicators are used to detect changes in environmental health, so Mudskippers can be considered as bioindicators
of marine pollution in Kuwait Bay, this needs further studies. [Nature and Science. 2010;8(5):79-89]. (ISSN: 1545-
0740).

Key words: Mudskippers, Intertidal Zone, Kuwait Bay

1. Introduction themselves up to 2 feet (60 cm) into the air (Piper,

Mudskippers are members of the subfamily 2007) .They have ability to breathe through their skin
Oxudercinae (tribe Periophthalmini, Murdy, 1989), and the lining of their mouth (the mucosa) and throat
within the family Gobiidae (Gobies). They are (the pharynx). This is only possible when the
completely amphibious fish that can use their pectoral mudskipper is wet, limiting mudskippers to humid
fins to "walk" on land (Swanson, and Gibb, 2004; habitats and requiring that they keep themselves moist.
Harris, 1960). Being amphibious, they are uniquely This mode of breathing, similar to that employed by
adapted to intertidal habitats, unlike most fish in such amphibians, is known as cutaneous air breathing
habitats which survive the retreat of the tide by hiding (Graham, 1997). Another important adaptation that aids
under wet seaweed or in tidal pools. Mudskippers are breathing while out of water is their enlarged gill
quite active when they are out of water, feeding and chambers, where they retain a bubble of air. These large
interacting with one another. Mudskippers constitute a gill chambers close tightly when the fish is above water,
group of 25 air-breathing species in four genera keeping the gills moist, and allowing them to function.
(Periophthalmodon, Periophthalmus, Boleophthalmus They supply oxygen for respiration also while on land
and Scartelaos) that are the most derived and the most (Graham, 1997). Digging of deep burrows in soft
amphibious of the ten genera of the teleost subfamily sediments that allow the fish to thermo regulate (Tytler
Oxudercinae (Gobiidae: Murdy, 1989; Clayton, 1993; and Vaughan, 1983), avoid marine predators during the
Graham, 1997; Aguilar, 2000). These fishes spend high tide when the fish and burrow are submerged
extensive periods of time out of water and have (Sasekumar et al.. 1994) and for laying their eggs
numerous physiological, morphological and behavioral (Brillet, 1969).
specializations for amphibious life (Gordon et al., 1969; Even when their burrow is submerged, mudskippers
Clayton, 1993; Graham, 1997; Lee and Graham, 2002). maintain an air pocket inside it, which allows them to
These fishes present a range of peculiar behavioural and breathe in conditions of very low oxygen concentration
physiological adaptations to an amphibious lifestyle. (Ishimatsu et al., 1998; Ishimatsu et al., 2000; Lee et al.,
These include: Anatomical and behavioural adaptations 2005). Eighteen species of the genus Periophthalmus
that allow them to move effectively on land as well as in have been described (Larson and Takita, 2004; Jaafar et
the water (Harris, 1960). As their name implies these al., 2009; Jaafar and Larson, 2008). Periophthalmus
fish use their fins to move around in a series of skips. argentilineatus grows to a length of about 9.5 cm and is
They can also flip their muscular body to catapult it feeds on small prey such as small crabs and other

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arthropods (Milward, 1974). Another species, and their eggs, which are deposited in the burrows, can
Periophthalmus barbarus, is the only oxudercine goby develop under severely hypoxic conditions. During the
that inhabits the coastal areas of western Africa (Murdy, mating season, the males become much more active,
1989). and their colors become more intense. Jaafar et al.
Mudskipper colonies are reported from the Bay of (2006) recorded the mudskipper, Periophthalmus
Kuwait in the Arabian Gulf area. Each fish digs his own walailakae from Singapore. This species most closely
deep burrow where it hides from disturbances and resembles Pn. schlosseri but with only one row of teeth
during the high tide. Under certain conditions, the single on the upper jaw, scales on the isthmus, and a different
fishes (most probably Boleophthalmus boddarti) build upper lip and jaw morphology. The authors (ibid) added
polygonal territories of a size of about one meter, that contrary to an earlier report, scales are present on
surrounded by dams, defended against rivals, and large the snout, intertidal, and isthmus of Ps. walailakae. The
enough to provide food (Microphytobenthos) (Höpner, two species can also be distinguished by size, external
1999). Often, the mudskipper form mixed colonies with morphology, and body color patterns. As stated by
digging crabs (Fiddler crabs -Ucaidae). Sayer and Harris (1961), these fishes present a range of peculiar
Davenport (1991) stated that amphibious behaviour in behavioral and physiological adaptations to an
fish has resulted in the colonization and eventual amphibious lifestyle. He (ibid) reported that these
domination by vertebrates of the terrestrial habitat. It include anatomical and behavioral adaptations that
is generally proposed that aquatic hypoxia, owing to allow them to move effectively on land as well as in the
metabolic oxygen consumption and organic decay, was water. Piper (2007) reported that they can also flip their
the most important selective force in the evolution of muscular body to jump up to 60 cm in the air. They
air-breathing vertebrates (Randall et al., 1981). There is have ability to breathe through their skin and the lining
evidently scope for detailed examination of emersion in of their mouth and throat .This is only possible when the
a number of amphibious fishes, testing a matrix of mudskipper body is wet, limiting mudskippers to humid
environmental and biotic stimuli, in an attempt to habitats and requiring that they keep themselves moist.
determine in more detail the reasons for such behaviour This mode of breathing, similar to that employed by
(Sayer and Davenport, 1991). Tytler and Vaughan (1983) amphibians, is known as "cutaneous air breathing"
reported that the annual range of body temperatures (Graham, 1997). Another important adaptation that aids
(14–35°C) of emergent mudskippers are substantially breathing while out of water is their enlarged gill
less than that of air temperatures (10–42°C) as a result chambers, where they retain a bubble of air. These large
of behavioural thermoregulation. Body temperatures gill chambers close tightly when the fish is above water,
generally match those of wet mud, which can be 7°C keeping the gills moist, and allowing them to function.
lower than air shade temperatures. They supply oxygen for respiration also while on land
Colombini et al. (1996) stated that activity patterns (Graham, 1997). As reported in the study of Tytler and
and zonation of the mudskippers were directly Vaughan (1983) diggings of deep burrows in soft
influenced by the synodic and tidal cycles and depended sediments that allow the fish to thermo- regulate avoid
more on environmental factors such as air temperature marine predators during the high tide when the fish and
and relative humidity than on the diel light cycle. burrow are submerged (Sasekumar, et al., 1994) and for
Chen et al. (2007) reported that the mudskipper, laying their eggs (Brillet, 1969). Periophthalmus
Boleophthalmus pectinirostris forms a territory during argentilineatus is one of the most widespread and
the cold season to keep a pool of water that encourages well-known species. It can be found in mangrove
diatom growth and enables the fish to engage in surface ecosystems and mudflats of East Africa and Madagascar
activity. Mudskippers have eyes at the top of the head east through the Sundarbans of Bengal, South East Asia
for an all-round view, while their mouth faces to Northern Australia, Southeast China and Southern
downwards to feed on the mud surface. Their pectoral Japan, up to Samoa and Tonga Islands .Another species,
fins are used like crutches to crawl over mud. Periophthalmus barbarus, is the only oxudercine goby
Mudskippers are a carnivorous opportunist feeder. that inhabits the coastal areas of Western Africa (Murdy,
Mudskippers dig deep burrows to escape predators and 1989). Both of these amphibious habits are completely
raise their young. They maintain an air pocket in their unsuited for normal fish tanks. Clayton (1987),
burrows to breathe. Even when their burrow is Clayton and Vaughan (1988) and Clayton and
submerged, mudskippers are seen to maintain an air Wright (1989) stated that the proximate mechanisms of
pocket inside it, which allows them to breathe in territorial behaviour in Boleophthalmus boddarti, an
conditions of very low oxygen concentration (Ishimatsu amphibious gobiid mudskipper that builds and
et al., 1998; Ishimatsu et al., 2000; Lee et al., 2005). maintains polygonal mud-walled territories provide a
Ishimatsu et al. (1998) reported that mudskipper fishes good example of the elastic disc concept of territories.
can maintain their metabolism while they are confined Clayton and Wright (1989) stated that at high
in mudflat burrows filled with oxygen-depleted water, population densities, the amphibious and herbivorous

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mudskipper Boleophthalmus boddarti construct mud their burrows and depositing them on their walls. As far
walls around their territories as a means of reducing as is known, this population in Sulaibikhat Bay is
aggression between neighbours. The authors (ibid) unique throughout the Indo-Pacific area (Clayton and
stated there were no significant differences in diatom Wells, 1994). Boleophthalmus stays always within its
density between territorial and non-territorial areas or mud walls, which enclose an area rather like a pasture
between grazed and non-grazed areas within territories. that has enough food for each individual to survive.
They concluded that the mud walls are considered to Inside each walled com- pound, Boleophthalmus digs a
play a secondary, indirect role in maintaining burrow up to one and a half metres deep. The burrow
populations of diatoms within territories. The mudflats may be anywhere in the territory, maybe right beside the
are subject to intense bioturbation, and that for the wall (Clayton and Wells, 1994).
Arabian Gulf area, mudskipper colonies are reported The mud is very good for burrowing in, since the
from the Bay of Kuwait. Each fish digs his own deep particles are very sticky, unlike sand. In all except the
burrow where it hides from disturbances and during very softest ooze, burrows will last for quite a long time.
high tide (Höpner, 1999). Under certain conditions, the Some burrows have a little chimney sticking up above
single fishes build polygonal territories of a size of the surface, others have small water-filled pools beside
about one meter, surrounded by dams, defended against the openings, and the reason for these variations is
rivals, and large enough to provide food unknown (Clayton and Wells, 1994). Boleophthalmus is
(Microphytobenthos). This scenery is rare even in very quick to dive into his burrow, and constantly alert
Kuwait Bay and difficult to be accessed; it is to danger. If they suspect a predator is around,
widespread on lower situated mudflats of the Khowre Boleophthalmus will merely raise his bulbous eyes
Musa area where it covers many km2. Even in the above the surface of the mud to check out the situation,
higher situated zone, mudskippers contribute decisively ducking down again at the slightest sign of trouble.
to the bioturbation power but do not form territories. Clayton and Wells (1994) reported that all
Often, they form mixed colonies with digging crabs, e.g. mudskippers are sexually monomorphic, which means
Fiddler crabs (Ucaidae). In Kuwait, Clayton and Wells that the males and females look exactly alike. However,
(1994) distinguished four main species of mudskippers during the breeding season, which extends from March
around the mudflats of Kuwait Bay and the Northern until late August, the males go through an elaborate
coast of Kuwait. The authors stated that the display that advertises their sex quite unmistakably.
mudskippers are also separated into zones. The After hatching, the larvae swim off with the tide and
mudskipper found on the high shore is called spend some time simply floating around with the other
Periophthalmus, a carnivorous mudskipper up to 15 types of plankton. Then they turn into very miniature
centimeters long which feeds on little crabs at low mudskippers and return to the mudflats where they
tide .Periophthalmus' eyes are well-adapted to vision in congregate in the soft mud areas. Here, as stated by
air, and when he's lying waiting for his prey only his Clayton and Wells (1994), they will remain until they
eyes stick up out of the muddy water. In order to keep reach maturity and manage to gain a territory of their
his eyes wet, Periophthalmus has little cups underneath own. Out of thousands of eggs, predators such as crabs
the eyes and when he blinks, the eyes roll down into his will take the vast majority of larvae and only a handful
skull and get remoistened by the water held in these will survive. The large xanthid crab, with its dark-tipped
little cups. claws, feeds on mudskipper larvae. Clayton and Wells
Slightly further down the shore the second species (1994) stated that the last two species of mudskippers
of mudskipper, called Boleophthalmus is the largest, found in Kuwait are both found much further out and so
measuring up to 25 cm and, like larger animals, is an are much harder to observe. They are also much less
herbivore. Boleophthalmus can be easily recognized by amphibious than their inshore cousins and spend their
their feeding action, a side-to-side head movement that time in permanently wet mud. The authors (ibid)
collects the fine surface film of diatoms and algae on stated that the larger of the two is called Scartelaos, and,
which they feed. Even in the hot summer, this species of although he is about as long as Boleophthalmus, is
mudskipper can remain active for several minutes at a much thinner. Clayton and Wells (1994) reported that
time out of water, and it can be recognized by its the final species of mudskipper is called Apocryptes and
characteristic mode of locomotion. The pelvic fins have is much smaller than any of the others. Since both
moved forward and fused together to form a little cup Scartelaos and Apocryptes inhabit areas of oozing wet
which he uses as a sort of crutch to balance on. His very mud, it is impossible for them to build burrows, so they
strong and well-muscled pectoral fins have moved down hide from predators by simply squirming down into the
the body and are, allowing him to swing along on them, soft mud out of sight.
using the life between the tides rear fin as a stabilizer. Clayton and Snowden (2000) observed the surface
Boleophthalmus build and maintain these polygonal activity of the carnivorous mudskipper Periophthalmus
walled territories by carrying mouthfuls of mud from waltoni Koumans 1941 was on mudflat in Sulaibikhat

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Bay, a muddy shore embayment in Kuwait Bay, at the coastal areas. A forestation of intertidal zones with
north-western head of the Arabian Gulf. They reported mangrove plants is considered a viable option to
that Each adult fish had a home range of between 2-3 improve coastal environment and enrich marine
m2 in which were located the main and subsidiary biodiversity. Mangrove plantations also protect the
burrows, the main one simply being the most frequently coastline from strong currents and support the
used. The burrows were of two types such that the accumulation of sediments and organic matter in the
entrance was either a double- turreted one (the ‘Y’ intertidal zones. These changes would improve the
shaped burrow) or was in a water-filled, saucer-like quality of mudflats and promote the survival and growth
depression. While some fish had a single burrow system of marine fauna (Sbandar et al., 20; Emabi, 1993; Ogino,
of either type, oth- utilise between two and six different 1993; Al Nafisi et a., 2009). The coastal zone of Kuwait,
ones. During the period of observation no other P. depending on the sediment nature and morphology, is
waltoni were seen to use them. classified into two main parts: Northern muddy
Mhaisen and Al-Maliki (1996) stated that the province and Southern rocky/sible for them to build
mudskipper Periophthalmus waltoni (Perciformes: burrows, so they hide from predators by simply
Gobiidae) in the Khor Al-Zubair estuary (Iraq) are squirming down into the soft mud out of sight.
infected with Myxobolus pfeifferi (Sporozoa), Clayton and Snowden (2000) observed the surface
Diplozoon sp. (Monogenea) and Neoechinorhynchus sp. activity of the carnivorous mudskipper Periophthalmus
(Acanthocephala). The State of Kuwait occupies waltoni Koumans 1941 was on mudflatal and growth of
approximately 17,800 km2 of the Northwestern part of marine fauna (Sbandar et al., 20; Emabi, 1993; Ogino,
the Arabian Gulf, between 28º30’ and 30º05’N, and 1993; Al Nafisi et a., 2009). The coastal zone of Kuwait,
46º33’ and 48º30’E. It comprises the mainland and nine depending on the sediment nature and morphology, is
offshore islands. Kuwait’s climate is characterized by classified into two main parts: Northern muddy
hot summers and mild winters. Temperature extremes province and Southern rocky/sandy part. These parts are
are high, with means during the warmest and coolest subdivided into several zones (Abou-Seida and
months ranging between 46.2ºC and 6.9ºC. Winter Al-Sarawi, 1990). The bay supports a thriving fishing
brings occasional frost. Rainfall is minimal, not industry and contains a site of an aquaculture facility of
exceeding 115 mm/year, but evaporation is very high, 80 net pen cages with a production of roughly 500 – 600
averaging 14.1 mm/d. The relative humidity is low, and metric tons ole and normal values for this region of the
strong, dry and hot, Northwesterly winds prevail during world. Kuwait Bay is an exposed area and stressed as a
summer, particularly in June and July (Al Nafisi et al., result of the extensive man-made activities, such as
2009). dredging, indiscriminate solid and liquid waste disposal
"Kuwait bay" is considered as one of the and over fishing. Al-Bakri et al., (1985) have concluded
characterized features of the Kuwaiti marine that the alteration of the coastal zone has resulted in
environment, which is an elliptically shaped bay that more impact to the ecosystem compared to the harsh
protrudes from the Arabian Gulf in Westward direction environmental condition. The environmental condition
at its Northwestern corner. Kuwait Bay is of a moderate of the coastal area became more critical as a result of
size (850 km²) with an average water depth of 5 m and a the war-related activities (Al-Ghadban et al., 1992).
maximum depth of 20 m at the entrance (Al-Ghadban, El-Sammak et al., (2005) compared the levels of
2004). The Northern shoreline is a pristine shore and in heavy metals in bottom sediments of Kuwait Bay and
contrast, the Southern part the bay hosts urban activities, Sulaibikhat embayment with the reported values of
major ports such as Shuwaikh and Doha Ports and three Dubai, Greek and Canada and highlighted higher values
major power and desalination plants (Doha East, Doha of cadmium and nickel. Al-Majed et al. (2004)
West and Subiya). The Bay of Kuwait presents a unique identified certain concentration of methyl mercury in
ecosystem and a significant nursery ground for many the embayment and attributed such concentration due to
fishes and shrimp species (Al-Yamani et al., 2004). the power plant and also due to the pervious industrial
Although biological and ecological data on the marine outfall and also due to the shipping activities and the
biota of the region is limited, with some coastal areas discharged into the area from the emergency outlets.
receiving more attention than others at least four critical Al-Ghadban et al. (1994) reported higher values of total
marine habitats, coral reefs, intertidal marshes, organic carbon content (more than 3%) in the area.
mangrove and sea grass beds, and kept forest, have been Coastal pollution has been increasing significantly over
recognized in the region (Price et. al., 1993). the recent years and found expanding environmental
In recent years, increased developmental activities problems in many developing countries. The discharges
and misuse of native vegetation have greatly degraded of industrial wastes have resulted in high metal
the coastal environment and marine ecosystems. In concentrations in the local marine environment,
addition, harsh weather conditions have accelerated the especially in the coastal sediments (Saad et al., 1981;
disappearance of vegetation cover in both inland and Mance, 1987; Ni et al., 2005). Al-Sarawi et al. (2002)

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reported high levels of metal discharges from power, Results and Discussion
thermal, desalination and water treatment plants and The Mudskippers observed in the present study appear
leakage from oil wells in Kuwait marine environment to be quite active when they are out of water, feeding
Glibert et al. (2004) stated that the bay supports a and interacting with each other (Fig. 2). The results of
thriving fishing industry and contains a site of an the present work are presented in the following items:
aquaculture facility of 80 net pen cages with a
production of roughly 500 – 600 metric tons of sea 1- Description and habitat
bream (Sparus auratus L.) per year. Mudskipper, In the present study, the mudskipper that are found
Periophthalmus waltoni is distributed in the northern in and around the mudflats of Kuwait Bay, are
region of Kuwait Bay’s tidal mudflats. They are the prey amphibious fishes that are called mudskippers. The
for many predators and thus, it is essential to evaluate results agree with those of Clayton and Wells (1994),
the bioaccumulation of metals toxicants in this fish. The who are separated the mudskippers into zones. The high
gills, skin and food are the main routes of accumulation shore mudskippers are called Periophthalmus, which
of metals by fish (Hein et al., 1993; Ni et al., 2005). The are carnivorous mudskippers up to 15 centimeters long
objectives of the present study were to study the and feeds on little crabs at low tide. The genus
ecological behavior of the mudskippers and to Periophthalmus is by far the most diverse and
determine the relationships between the gobies fish widespread genus of mudskippers that contains eighteen
behavior and the Kuwait Bay’s mudflats habitat. described species (Larson and Takita, 2004).
They are uniquely adapted to intertidal habitats,
2. Materials and methods unlike most fish in such habitats, which survive the
The studied Mudskippers in the present study are retreat of the tide by hiding in tidal pools. Mudskippers
monitored along the Kuwait Bay shores (Fig. 1) during need to live in habitats that are hot and humid in order
the very hot summer season, near delivery hospital, to breathe, where air and water temperature range from
Shuwaikh area, Kuwait Governorate. They are 75 to 86 °F and humidity from 60 to 80 %. Mudskippers
particularly abundant in the muddy shore. However, a are only active when the temperature is above 55 °F.
number of fifty mudskipper samples are collected This result agrees with the results of Clayton and Well
randomly to be examined for parasites. (1994).
Other adaptation of mudskippers to life on land is
Field observations: their huge goggly eyes at the top of their heads for an
The procedure followed for monitoring the mud all-round view, while their mouth faces downwards to
skippers is described by Clayton and Wells (1994), that feed on the mud surface. These eyes sit on stalks while
comprises using a sheet of plywood or a big stone to the rest of their bodies remain safely underwater.
help to distribute our weights more evenly across the Unlike other fishes, mudskippers prefer to swim with
mud, as a surface crust of dried mud may seem quite their heads above water, their eyes giving them a good
hard, but if it cracks one, and one will sink right down 360-degree view. To keep their eyes moist when they
into the soft ooze below. But if one chooses the right are on land, the eyes can be retracted to dip them into
type of shore he should be able to observe the water that collects at the bottom of the eye socket.
mudskippers from the safety of dry land. Fifty Mudskippers are probably the only fish with movable
specimens of the amphibious gobies (2.17œ5.20 cm eyelids (Fig. 3). The pectoral fins of the mudskippers
standard length (SL); 0.16œ2.70 g body wet weight) are used like crutches to crawl over mud.
were collected using hand-nets during low tides in
Kuwait Bay. Species are abundant and successfully
surviving in the mud flat pool during low tides. The
caught specimens were placed in small plastic
containers, and kept alive in aerated sea-water prior to
further studies in the College laboratory. Taxonomic
identification and confirmation of the species were
based on description given in Clayton and Wells (1994).
Amphibious goby was identified as Periophthalmus sp.

Laboratory observations:
Collected specimens were examined for infestation
with external parasites, and then dissected for detecting
internal parasites in the body cavity and all other organs.

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2- Territorial behaviour
Mudskippers are noticed to dig deep burrows to
escape predators and raise their young. They maintain
an air pocket in their burrows to breathe. Chen et al.
(2007) reported that the mudskipper, Boleophthalmus
pectinirostris, maintained territories in farming ponds
during the cold season between November and
February. Two types of territory are found, one is
surrounded by mud-walls, and the other is without
mud-walls. Both types of territory are entirely covered
by shallow water (Fig. 4).
The area of walled territories was significantly bigger
than that of non-walled ones. The nearest neighbour's
distance of walled territories was significantly shorter
than that of non-walled territories. This description is
detected in the present study.
Fish surface activity occurred between 1200 and
1500 hours at which time the territorial water
temperature reached a maximum, being significantly
higher than that of the air, mud surface or deep burrow.
The territorial sediments exhibited a significantly
greater benthic micro- algal biomass as compared to the
non-territorial sediments. The present study indicates
that B. pectinirostris forms a territory a pool of water
Figs. 2A &2B. The observed Mudskippers appear to that encourages diatom growth and enables the fish to
be quite active when they are out of water. engage in surface activity (Fig. 4).
Clayton (1987), Clayton and Vaughan (1988) and
Clayton and Wright (1989) stated that the proximate
mechanisms of territorial behaviour in Boleophthalmus
boddarti, an amphibious gobiid mudskipper that builds
and maintains polygonal mud-walled territories
provides a good example of the elastic disc concept of
territories. The occurrence of wall building is density
dependent and a contiguous mosaic of territories is only
produced at high fish densities. Wall removal and
replacement experiments show that the mud-wall acts as
a visual barrier and reduces aggression between
neighboring territorial fish.
Clayton and Wright (1989) stated that at high
Fig. 4. Walled territories. population densities, the amphibious and herbivorous
mudskipper Boleophthalmus boddarti construct mud
walls around their territories as a means of reducing
aggression between neighbours. Because of the walls,
territories contain pools of water and exposed mud
slopes. Whilst the density of benthic diatom prey was
highly variable, the highest was found on the exposed
mud slopes and the lowest on the boundary walls. Fish
grazed mainly on the mud slopes. No significant
differences were found in diatom density between
territorial and non-territorial areas or between grazed
and non-grazed areas within territories. The variation of
the diatom density, however, was reduced within
territories. The mud walls play an indirect role in
Fig. 3. Mudskippers are probably the only fish with maintaining populations of diatoms within territories.
huge eyes for an all-over view. Höpner (1999) reported that the mudflats are
subject to intense bioturbation, and that for the

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Arabian Gulf area, mudskipper colonies are reported empty lot.

from the Bay of Kuwait. Each fish digs his own deep Naturally, the owners do not take kindly to this and
burrow where it hides from disturbances and during will chase the intruder out, into a neighboring territory,
high tide. The author (ibid) stated that under certain where upon the neighbours will take up the pursuit and
conditions, the single fishes (most probably a splendid chain reaction of indignant outrage will
Boleophthalmus boddarti) build polygonal territories of follow the poor trespasser all the way back to the soft
a size of about one meter, surrounded by dams, muddy area where belongs.. The main factor in ensuring
defended against rivals, and large enough to provide a constant supply of vacant territories is predation and
food(microphytobenthos). Clayton and Wells (1994) this is especially prevalent during the spring when the
stated that while this scenery is rare even in Kuwait Bay weather is cold and the fish are rather sluggish as a
and difficult to be accessed, it is widespread on lower result. Shore birds are their worst enemies.
situated mudflats of the Khowr-e Musa area where it Clayton and Wells (1994) stated that most of the
covers many km2. Even in the higher situated zone, fish found in the intertidal zone come there at high tide
mudskippers contribute decisively to the bioturbation to feed. However, to see the fish, which have best
power but do not form territories. Often, they form adapted to the amphibious life between the tides we
mixed colonies with digging crabs (Ucaidae). This have to go to the third main coastal region of Kuwait:
result agrees with the results of the present study. the mudflats. Even when their burrow is submerged,
Boleophthalmus build and maintain these polygonal mudskippers are seen to maintain an air pocket inside it,
walled territories by carrying mouthfuls of mud from which allows them to breathe in conditions of very low
their burrows and depositing them on their walls. As far oxygen concentration. This result accommodates with
as is known, this population in Sulaibikhat Bay is the results of (Ishimatsu et al., 1998; Ishimatsu et al.,
unique throughout the Indo-Pacific area (Clayton and 2000; Lee et al., 2005).
Wells, 1994)! Clayton and Wells (1994) observed mudskippers dig
Although they are found all over Kuwait Bay, the deep burrows to escape predators and raise their young
most interesting population of Boleophthalmus can be and that they maintain an air pocket in their burrows to
seen in Sulaibikhat Bay where they construct and breathe. During the mating season, the male's colors
maintain and elaborate and continuous network of become more intense, and they become much more
polygonal walled territories (see Fig. 3 and 4). Most of active. They (ibid) stated that males leap and flip in the
these territories are five-sided with an average area of air and even stand on their tails, all to attract the
1.69 square metres. The ones with only four sides tend females. The male becomes more aggressive forward
to be smaller, while those with six or seven sides are a other males with biting and raising its dorsal fins. If the
little bigger. These territories are unusual in that each is female is attracted to the male, she does her own mating
occupied by a single fish of either sex. ritual with distinctive movements. She then goes into
Boleophthalmus stays always within its mud walls the male's burrow where she lays her eggs in a special
which enclose an area that has enough food for each part of the burrow the male builds her. The males then
individual to survive. They build the walls with mud fertilize the eggs and takes over responsibility for the
from their burrows which they carry up in their mouths eggs. The above descriptions are noticed in the present
and spit out onto the section in need of repair. If part of study. These fishes present a range of peculiar
the wall is destroyed, they will work hard to repair it behavioral and physiological adaptations to an
within a matter of hours during low tide. The best time amphibious lifestyle. He reported that these include:
to see these walled territories is at low tide when the sun Anatomical and behavioral adaptations that allow them
is low in the sky. If there is a mild winter, some of the to move effectively on land as well as in the water
walls will last for more than a year .Inside each walled (Harris, 1961). They can also flip their muscular body to
compound, Boleophthalmus digs a burrow up to one and catapult themselves up to 60 cm into the air. This
a half metres deep. The burrow may be anywhere in the observation is noticed in the present study (Piper, 2007).
territory, maybe right beside the wall (Clayton and Another important adaptation that helps the
Wells, 1994). mudskippers to breath while they are out of water is
Young Boleophthalmus don't have their own their gill chambers, where they keep a bubble of air.
territories, due to lack of suitable space. Instead, they These large gill chambers close tightly when the
are forced to live in areas where the mud is too soft for mudskipper is above the water, allowing the gills to be
either walls or burrows, burying themselves in soft ooze kept moist, and make them work. They supply oxygen
whenever a predator threatens. Obviously, this is not as for respiration also while the gobies are on the land
safe as a nice deep burrow, so the young mudskippers (Graham, 1997).
are live between the tides constantly on the lookout for As reported in the study of Tytler and Vaughan
a vacant territory and constantly trespass both in order (1983), digging of deep burrows in soft sediments that
to feed and to investigate the chances of finding an allow the fish to thermoregulate , avoid marine

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 Nature and Science 2010;8(5)

predators during the high tide when the fish and burrow then arches his back. It has two sets of paired fins,
are submerged (Sasekumar et al., 1994) and for laying pectorals near his chest, and pelvic fins at the hips. The
their eggs (Brillet,1969). pelvic fins have moved forward and fused together to
Tytler and Vaughan (1983) stated that in winter, form a little cup, which is used as a sort of crutch to
mudskippers avoid low surface temperatures by balance on. This description agrees with the results of
remaining in their burrows, in the present study, no the present study.
mudskippers were detected during winter months. In Periophthalmus, was observed sitting on the
summer, body temperatures are kept lower than ambient mudflats splashing water over his body with his pectoral
by selecting areas where evaporative cooling is high. fins for about several minutes before moving back to his
Body temperatures generally match those of wet mud, burrow. This agrees with the description of Clayton and
which can be 7°C lower than air shade temperatures. Wells (1994). These burrows are quite remarkable; they
Chen et al. (2007) stated that the mudskipper are y -shaped, with two entrances and are more than a
Boleophthalmus pectinirostris maintained territories in meter deep in order to get down to permanent water
farming ponds during the cold season between level. The mud is suitable habitat for burrowing in,
November and February. since the particles are very sticky, unlike sand. In all
The present study results accommodate with the except the very softest ooze, burrows will last for quite
above observations. a long time (Clayton and Wells, 1994).

3- Feeding
Periophthalmus needs to be out of water to feed
and it needs to keep his skin wet, it can absorbs oxygen
through its skin as well as through its gills. Some times,
it will fill its mouth with water to help him to survive
for a longer time out side the water (Fig. 6), since the
inside of its mouth has lots of tiny blood vessels that
absorb oxygen. However, this water isn't 'essential; the
mudskipper loses it anyway when it eats his prey
(Clayton and Wells, 1994).
Ishimatsu et al., (1998) reported that mudskipper
fishes can maintain their metabolism while they are
confined in mudflat burrows filled with oxygen-
depleted water, and their eggs, deposited in the burrows,
can develop under severely hypoxic conditions.
Their feeding action, a side-to-side head
movement that collects the fine surface film of diatoms
Fig. 5. Periophthalmus' eyes are well-adapted to see in and algae on which they feed can easily recognize
air and out of the muddy water. Boleophthalmus. Terns are another ménage to the
mudskipper that they catch by flying along about three
Periophthalmus' eyes are well-adapted to see in air, to five metres up, then dive-bombing their prey with
and when they are lying in wait for their preys only their bayonet-like beaks. Sea gulls use a craftier
their eyes stick up out of the muddy water (Fig. 5). In approach, skimming very low, just above the mud walls
order to keep their eyes wet, Periophthalmus have and trying to catch the Boleophthalmus unawares.
little cups underneath their eyes and when they But, Boleophthalmus like all the other
blinks, the eyes roll down into their skull and get mudskippers, is very quick to dive into his burrow, and
remoistened by the water held in these little cups. Even constantly alert to danger. If they suspect a predator is
in the heat of summer, this species of mudskipper can around, Boleophthalmus will merely raise his bulbous
remain active for several minutes at a time out of water. eyes above the surface of the mud to check out the
They are seen periodically rolling on their sides to help situation.
to keep their skins wet.
Every five minutes or so, Periophthalmus will dash 4- Infection of the mudskippers with parasites:
back to its burrow to cool off and refresh itself. This External examination and dissection of 50 samples
species of mudskipper can be recognized by its of the mudskipper collected from the studied area
characteristic way of locomotion. It lifts his tail, and indicates no parasitic infections.

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 Nature and Science 2010;8(5)

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