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A review of the biology and ecology of the whale shark



The information available on the biology and ecology of the whale shark is reviewed, and is updated from material published since 1986. Research work carried out on the seasonal aggregation of whale sharks at the Ningaloo Reef in Western Australia is summarized. Future research studies on whale sharks in the Ningaloo Marine Park are discussed in the context of management of sustainable whale shark interaction tourism.
Journal of Fish Biology (1997) 51, 1219–1234
A review of the biology and ecology of the whale shark
J. G. C
Marine Conservation Branch, Department of Conservation and Land Management,
47 Henry Street, Fremantle, Western Australia 6160, Australia
(Received 11 January 1997, Accepted 4 July 1997)
The information available on the biology and ecology of the whale shark is reviewed, and is
updated from material published since 1986. Research work carried out on the seasonal
aggregation of whale sharks at the Ningaloo Reef in Western Australia is summarized. Future
research studies on whale sharks in the Ningaloo Marine Park are discussed in the context of
management of sustainable whale shark interaction tourism.
?1997 The Fisheries Society of the British Isles?1997 The Fisheries Society of the British Isles
Key words: whale shark; Rhiniodon typus; Ningaloo Reef.
The whale shark, Rhiniodon typus Smith 1828, has a very widespread distri-
bution and occurs throughout the world’s tropical and warm temperate seas, and
yet knowledge of its biology and ecology is very limited. Like many other shark
species, the species has innate biological characteristics, such as large size, slow
growth, late maturation and extended longevity, that probably limit recruitment
and make it particularly susceptible to exploitation. These characteristics may
also mean that populations are slow to recover from any overexploitation (Jones
& Kaly, 1995).
With the worldwide growth of marine nature-based tourism, recreational
snorkelling and scuba diving, there has been a steady increase in the number of
encounters with whale sharks. In a few locations, such as the Ningaloo Reef in
Western Australia, the Galapagos Islands, the islands of the Andaman Sea o
the west coast of Thailand, and the Sea of Cortez and Baja California in the
eastern Pacific, where occurrences of whale sharks appear to be predictable, they
are being targeted increasingly by commercial tourist operations. These opera-
tions provide a rare opportunity for close encounters between humans and large
marine fauna, but may result in unknown eects on the shark’s behaviour and
ecology. International conservation status of the species is unclear—it is listed as
having an ‘ indeterminate ’ status on the World Conservation Union’s Red List
of Threatened Animals (IUCN, 1994). This category applies to animals known to
be ‘ endangered ’, ‘ vulnerable ’ or ‘ rare ’, but there is not enough information
available to say which of these three categories is appropriate. Casey et al. (1992)
considered the whale shark to be at potential risk from pelagic fisheries. There
are indications that even small traditional fisheries may be unsustainable, with
catches from a seasonal fishery in the Philippines declining over recent years, but
the reason for this downward trend is unknown (Trono, 1996). Globally,
commercial fisheries for whale sharks are limited at present, but may expand
from an increased demand for food products.
0022–1112/97/061219+ 16 $25.00/0/jb970526 ?1997 The Fisheries Society of the British Isles
A seasonal aggregation of whale sharks occurs in the waters of the Ningaloo
Marine Park in Western Australia (Fig. 1) from March to May each year. Over
the last 4 years, this predictable occurrence has led to the development of a small
but expanding tourist industry, focusing on human–whale shark interactions.
From 1993 onwards, commercial whale shark tourism has been managed by the
F. 1. Ningaloo Marine Park. ., Cape Range National Park; , Ningaloo Marine Park (Common-
wealth jurisdiction); , Ningaloo Marine Park (State jurisdiction); /, Sanctuary zones.
1220 ..
Western Australian Department of Conservation and Land Management
(CALM) through a system of controls, including the licensing of a limited
number of operators for whale shark interaction tours within the Marine Park.
Presently, it is unclear whether increased tourism pressure is generating any
short- or long-term detrimental impacts on individuals sharks or the group as a
whole. The natural variability in whale shark abundance and distribution, the
reasons for the aggregation at the Ningaloo Reef and the carrying capacity of the
industry are all unknown. Consequently, evidence of any impacts is dicult
to obtain and interpret. With the limited information currently available a
precautionary approach to management has been adopted.
To improve the management of the interaction, and in the long-term, to
provide the scientific basis to determine if current management strategies need to
be modified to minimize any impacts, CALM has developed a management
programme for whale shark interaction tourism. This has included an extensive
and up-to-date review of the biology and ecology of the species, in the context of
the seasonal aggregations at the Ningaloo Reef. Once more information is
available and appropriate monitoring programmes are implemented it will be
possible to ensure that the whale shark population in the Ningaloo Marine Park
is not being subjected to an unacceptable level of disturbance, and that
development of whale shark tourism is sustainable and equitable.
The published literature on whale sharks is extensive and consists mainly of
sightings records, anecdotal reports, speculative reviews of distribution and
movement patterns, and limited observations of general biology, feeding and
behaviour. Much of the material is derivative, superficial in content, and of
limited scientific use. The definitive bibliography (Wolfson & Notarbartolo di
Sciara, 1981) collated all available literature to 1980, listed and annotated 345
references, and categorized these under 19 broad headings. This review was
updated later by a consolidation of reported sightings (Wolfson, 1986). Silas
(1986) updated records from Indian waters and carried out a comprehensive
review of the information available on the whale shark’s biology and ecology.
The aim of this overview is to examine these and other key papers published
before 1986, plus any further material published since 1986, and to review
research carried out at Ningaloo Reef in the last 10 years, in order to update the
existing knowledge of the biology and ecology of R. typus. A comprehensive
reference database and reprint collection have been established at CALM’s
Marine Conservation Branch. Currently, the database lists 541 references and
contains 130 articles which were either not listed in previous reviews, or have
been published since 1986. The relational database and reprint collection are
being made available to researchers carrying out studies on whale sharks.
The species was first described and named by Dr Andrew Smith from a
specimen harpooned in Table Bay, South Africa in 1828 (Smith, 1828,1829,
1849). Historically, there has been considerable synonymy at family, generic
and specific level, and in 1984 the International Commission on Zoological
Nomenclature suppressed previous generic variations in favour of genus
Rhincodon, family Rhincodontidae (Melville, 1984). Alternate generic names
       1221
most commonly used are Rhiniodon and Rhineodon. Systematically, Rhinco-
dontidae (with the single type species Rhiniodon typus) is placed in the order
Orectolobiformes, which also includes nurse sharks (Ginglymostomatidae),
leopard sharks (Stegostomatidae), and wobbegongs (Orectolobidae). The inter-
relationships between these families are based upon a number of anatomical and
morphological similarities, including skeletal anatomy, tooth and dermal
denticle morphology, fin placement and barbel morphology (Compagno, 1973;
Dingerkus, 1985).
The whale shark is the world’s largest living fish. It is characterized externally
by a broad, flattened head, a very large and nearly terminal mouth, very large gill
slits, three prominent longitudinal ridges on its upper flanks, a large first dorsal
fin, a semi-lunate caudal fin and a unique ‘ checkerboard ’ pattern of light spots
and stripes on a dark background (Compagno, 1984;Last & Stevens, 1994). The
function of this distinctive pattern of body marking is unknown. Many
bottom-dwelling sharks have bold and disruptive body markings that act as
camouflage through disruptive coloration (Bass, 1978). The whale shark’s
markings could be a result of its phyletic relationship with bottom-dwelling
orectolobiform carpet sharks. Sharks have a high degree of visual development
(Gruber, 1977) and distinctive markings in a pelagic species could be linked to
social activities such as postural displays and recognition processes (Myrberg,
1991). Another possibility is that these pigment patterns could be an adaptation
for radiation shielding, important in a species that may spend a significant
proportion of time in surface waters possibly exposed to high levels of ultraviolet
The whale shark is one of three species of very large filter-feeding sharks that
occur in Western Australian waters, the other two being the basking shark
Cetorhinus maximus (Gunnerus), and the megamouth shark Megachasma
pelagios (Taylor, Compagno & Struhsaker).
R. typus is thought to be cosmopolitan in distribution, occurring in all tropical
and warm temperate seas apart from the Mediterranean. It is found in a band
around the equator between about 30)N and 35)S, in both coastal and oceanic
waters (Compagno, 1984). It occurs throughout the Indian Ocean and has been
reported from the Maldives, Seychelles and Comores Islands, and along the
coastlines of Madagascar, South Africa, Mozambique, Kenya, Pakistan, India,
Sri Lanka, Thailand, Malaysia and Indonesia. In Australia, whale sharks occur
mainly onorthern Western Australia at the Ningaloo Reef, the Northern
Territory and Queensland, with isolated reports from New South Wales and
Victoria (Wolfson, 1986).
In contrast to the majority of orectolobiform sharks, which are benthic species,
the whale shark has a pelagic habitat. Iwasaki (1970) collated and analysed
environmental data from skipjack tuna fishing vessels that encountered whale
sharks in the western Pacific Ocean from 1955 to 1967. Analysis of air
temperatures, seawater temperature and salinity profiles, and wind and current
patterns revealed changes in the seasonal frequency and distribution of whale
1222 ..
sharks that appeared to be linked to a number of environmental variables,
including the warm Kuroshio current and warm SSW–WSW winds. Othe east
coast of Japan their range encompassed areas with surface water temperatures
from 18 to 30)C, but they appeared to prefer locations with surface water
temperatures between 21–25)C, where cool nutrient-rich upwellings mingle with
warm surface waters of salinities between 34–34.5‰ (Iwasaki, 1970). These
conditions may well be optimal for the production of the planktonic and
nektonic prey upon which the sharks feed.
Sightings of whale sharks, made during aerial surveys along the south coast of
Texas, occurred in waters with surface temperatures of 29)C(Homan et al.,
1981). Arnbom & Papastavrou (1988) reported several sightings of whale sharks
in the Galapagos Islands, in an area of very deep water (2000–3000 m) close to
the edge of the continental shelf, known for its upwellings and high primary
productivity. Surface seawater temperatures recorded during these encounters
were between 23·5 and 26·5)C.
The whale shark is a filter-feeder that appears to feed on a wide variety
of planktonic and nektonic prey, including small crustaceans such as krill,
crab larvae and copepods, small schooling fishes such as sardines, anchovies,
mackerel, and occasionally larger prey such as small tuna, albacore and squid
(Compagno, 1984;Last & Stevens, 1994). Also, phytoplankton and macroalgae
may form a component of the diet (McCann, 1954;Kaikini et al., 1959;
Satyanarayana Rao, 1986;Karbhari & Josekutty, 1986). An analysis of the
stomach contents of a specimen caught othe coast of India in 1961 revealed a
variety of material, ‘ including large quantities of zooplankton, partly digested
remains of fish, crustaceans, molluscs, and small quantities of seaweed and algae,
undoubtedly suggesting an omnivorous diet ’ (Silas & Rajagopalan, 1963).
However, it is possible that algal matter found in the gut contents of these
specimens could have been swallowed accidentally, either during the course of
normal feeding activities or whilst being captured.
The animal is not dependent on forward motion to operate its filtration
mechanism, but rather relies on a versatile suction filter-feeding method, which
enables it to draw water into the mouth at higher velocities than dynamic
filter-feeders, such as the basking shark (Compagno, 1984). This enables it to
capture larger, more active nektonic prey as well as zooplankton aggregations,
but probably means it can filter a far smaller volume of water making it less
ecient in concentrating diuse planktonic food. Therefore, the whale shark
may be more dependent on dense aggregations of prey organisms. Taylor et al.
(1983) reviewed the feeding biology and filter apparatus of R. typus in relation to
M. pelagios and C. maximus and concluded that the dense filter screens of the
former act as more ecient filters for short suction intakes, in contrast to the
flow through systems of the latter two species.
Individuals have been observed feeding passively (cruising with mouth agape)
and also sometimes hanging vertically in the water and feeding actively by
opening their mouths and sucking in prey-rich water. They have been seen
feeding in this manner on aggregations of small crustaceans (including
euphausids), squid, anchovy and sardines (Silas, 1986). They have also been
       1223
observed coughing, which is thought to be a mechanism employed to clear or
flush the gill rakers of accumulated food particles. Groups of individuals have
been observed feeding actively at dusk or after dark by ploughing through the
surface waters with mouth agape and jaw distended, sometimes also moving
their heads from side to side vacuuming in sea water rich in prey, or aggressively
cutting swathes through schools of prey (Clark, 1992;Taylor, 1994). At the
Ningaloo Reef in Western Australia, individuals and groups of whale sharks
have been observed feeding actively on swarms of the tropical krill
Pseudeuphausia latifrons (G. O. Sars) (Taylor, 1994).
Information about the reproduction and development of the whale shark is
very limited. Historically, there has been much debate about their mode of
development, and it was unclear whether the whale shark is oviparous (egg cases
expelled from the female’s body and hatching on the sea floor) or ovo-viviparous
(egg cases hatching in utero, with the female giving birth to live young).
Southwell (1912/1913) reported that a specimen taken othe coast of Sri Lanka
was found to contain a ‘ very large ovary, oviduct full of eggs, 16 cases counted,
same form as in dogfish ’. This observation suggested oviparity. Until recently,
the only known whale shark embryo was a near-term 36-cm specimen retrieved
alive from a large egg case trawled from the sea floor in the Gulf of Mexico in
1953 (Baughman, 1955;Reid, 1957;Garrick, 1964). Whilst this appeared to
suggest an oviparous mode of development, Wolfson (1983) believed it was more
likely that the egg case had been aborted, and that the whale shark is actually
ovo-viviparous. This conclusion was based on the absence of other occurrences
of ‘ free-living ’ egg cases and the fact that the Gulf of Mexico egg case was
extremely thin and lacked tendrils (which anchor the egg case to the sea floor).
The embryo also had large reserves of yolk and only partially developed gill
sieves, which combined with the discovery of umbilical scars on a free-living
juvenile of 55 cm length (Wolfson, 1983), provide further support for the theory
that whale sharks have an ovo-viviparous mode of development.
In July 1995, a female whale shark, measuring approximately 11 m in length,
was harpooned othe eastern coast of Taiwan (Joung et al., 1996). The twin
uteri of this specimen were found to contain 300 embryos, from 42 to 63 cm in
length. Fifteen of the embryos were alive and one, measuring 58 cm in length,
was reared for 143 days in an aquarium in Japan where it developed from fetal
through to juvenile stage (Leu et al., unpublished). This discovery finally
confirms that the species is a live-bearer, with an ovo-viviparous mode of
A total of only nine juveniles, ranging from 55 to 93 cm in length, has been
recorded in the literature (Wolfson, 1983;Anon., 1989;Kukuyev, 1996). One of
the juveniles recorded by Kukuyev (1996) was found in the stomach of a blue
shark Prionace glauca (L.), caught in central tropical Atlantic waters. A further
specimen, measuring 61 cm in length, was found in the gut contents of a blue
marlin Makaira mazara (Jordan & Snyder) caught othe northern coast of
Mauritius in 1993 (D. Goorah, pers. comm.). The juvenile was alive when
1224 ..
recovered from the marlin’s stomach, as it had just been ingested prior to the
intake of a bonito Sarda orientalis (Temminck & Schlegel).
There are no confirmed records of whale sharks between 93 cm and 3 m.
Animals over 3 m in length are encountered worldwide. Most specimens
reported in the literature are between 4 and 10 m, but maximum total length is
uncertain. The largest accurately measured specimen was 12 m in length
(Karbhari & Josekutty, 1986), but there is a report of a specimen from the
Seychelles that measured nearly 14 m in length (Wright, 1870), and a specimen
measuring just over 14 m was landed in India in 1975 (Devadoss et al., 1990).
Whale sharks may reach possibly as much as 18 m in length, although the very
large specimen reported from the Gulf of Siam (Smith, 1925) was not measured
accurately and therefore total length may have been overestimated.
Information about size at sexual maturity and longevity is sparse. Taylor
(1994) speculated that whale sharks do not reach sexual maturity until they are
over 30 years of age, and that they may have a life span of over 100 years. Also,
he observed that the only sexually mature male seen at the Ningaloo Reef was
probably over 9 m in length. Two large female sharks captured othe Indian
coastline (Pai et al., 1983;Satyanarayana Rao, 1986), which measured 8–9 m in
length, were both found to have immature ovaries. The evidence suggests that
sexual maturity in both sexes may not occur until the sharks are over 9 m in
length. Data on sex ratio is also very limited. Of 31 specimens reported from
India, 17 were male and 14 were female (Silas, 1986). Taylor (1994) observed
that the majority of whale sharks encountered at Ningaloo are immature males.
On the basis of current information it is not possible to say whether sexual
segregation, of either a behavioural or geographical nature, occurs.
Virtually nothing is known about growth rates or ageing in whale sharks. As
accurate measurements of size parameters are dicult to obtain from whale
sharks in the water, morphometric studies of this species have made little
progress. Generally, growth curves for sharks have been derived from age
estimates based upon growth zones (bands or rings) in calcified structures such as
vertebral centra. The growth of whale sharks has been studied opportunistically
from animals held in captivity in the Okinawa Expo Aquarium in Japan (Uchida
et al., 1990). One of these animals was fed food laced with tetracycline, on
several occasions over a period of more than a year (Cailliet et al., 1986).
Preliminary results indicated that one pair of growth zones was deposited per
year in captivity. How this relates to growth in the wild can be determined only
by examination of growth zones in vertebral centra samples collected from dead
Studies of this nature are hampered by sample size, as very few dead animals
are available for collection of hard tissues. There are no records of whale shark
strandings along the Western Australian coastline, although there is an uncon-
firmed report of a small (2 m length) whale shark being found on the beach at
Sandy Bay, North West Cape (Fig. 1) in 1982 (N. Nannup, pers. comm.). For
the last 3 years, researchers in South Africa have been retaining vertebrae
samples from specimens that strand occasionally along the KwaZulu/Natal coast
       1225
(G. Cli, pers. comm.). Some information on ageing of whale sharks may be
available from subsequent examination of these samples.
In general, occurrences of whale sharks appear to be sporadic and unpredict-
able, which is partly a reflection of the lack of knowledge about the animal’s
habitat and ecology. Generally, whale sharks are encountered singly but
aggregations of over 100 animals have been seen (Anon., 1961), which suggests
that schooling activity does occur. They are usually observed on or near the
surface and at times have been seen apparently basking.
Whale sharks are thought to be highly migratory but currently there is no
direct evidence to support this. Their movements are related probably to
increases in local productivity such as plankton blooms and invertebrate
spawning events, with associated increases in zooplankton and bait fish shoals,
and also to changes in water temperature, currents, winds and other environ-
mental parameters (Compagno, 1984). It has been suggested that whale sharks,
with their suction filter-feeding strategy, are probably more dependent on
localized productivity events (J. Stevens, pers. comm.). Dierent locations
appear to be preferred at various times of the year and they may undertake either
fairly localized migrations or alternatively large-scale transoceanic movements,
governed by the timing and location of production pulses and possibly by
breeding behaviour. Seasonal migrations have been postulated for various areas
but more information is needed to confirm these patterns (Wolfson, 1986).
Taylor (1994) postulated that the appearance of whale sharks at the Ningaloo
Reef during the austral autumn period is linked to the high levels of productivity
associated with mass synchronous coral spawning events after the March and
April full moons (Simpson, 1991). Currently, it is unknown whether the whale
sharks present at Ningaloo in March to May are resident in the eastern Indian
Ocean throughout the rest of the year. These animals could move oshore to
deeper waters and may be seen only when they come inshore and into surface
waters to exploit periodic increases in plankton productivity, such as those
around the time of mass coral spawning. Inshore sightings of significant
numbers of whale sharks have been made in December and January along the
Western Australian coastline between Kalbarri and Shark Bay (P. Wieland, pers.
comm.). Unconfirmed reports suggest the occurrence of numbers of whale
sharks around the Montebello Islands on the North West Shelf of Western
Australia at the same time of year. Whether either of these separate aggregations
consist of the same individuals as those aggregating at the Ningaloo Reef during
the autumn months is not known at present.
Episodic aggregations of whale sharks also occur in two other locations in
Australian waters. During December and January significant numbers of whale
sharks have been reported from Christmas Island in the Indian Ocean
(J. Stevens, pers. comm.) and this aggregation occurs at the same time that the
red crab Gecarcoidea natalis (Pocock) spawns en masse. In the Coral Sea regular
sightings of whale sharks occur in October and November in association with
aggregations of tuna Thunnus spp. (Gunn et al., 1992). It is believed that these
aggregations may be associated with large concentrations of spawning lantern
fish Diaphus spp., rather than with synchronous coral spawning along the Great
1226 ..
Barrier Reef that occurs during the same period (J. Stevens, pers. comm.). In the
Maldives, whale sharks show distinct seasonal movements, in phase with the
changing monsoons and associated current movements, as well as up-welling
events and plankton blooms (Anderson & Ahmed, 1993).
There are several reports in the literature of seasonal aggregations of whale
sharks in Indian coastal waters from December to April (Silas, 1986), in the
Seychelles during August and November (R. Salm, pers. comm.), and othe
west coast of Mexico from Cabo San Lucas to Acapulco from March to August
(Wolfson, 1986,1987). Significant numbers of whale sharks have been reported
from inshore waters osouthern Mozambique and the northern coast of South
Africa, from October through to March (A. Giord, pers. comm.). Since 1993,
researchers from the Shark Research Institute in South Africa have been
carrying out whale shark aerial surveys and a tagging programme along the
KwaZulu/Natal coastline (A. Giord, pers. comm.). An aerial survey in
January 1994 recorded a total of 95 whale sharks along 110 km of coastline.
During the 1994 and 1995 seasons, a total of 72 sharks were tagged. The
researchers are hoping to expand their tagging eort along the east African
coast, using tags of identical design but with dierent colour coding according to
the tagging location. Returns could assist in the determination of large-scale
migratory patterns throughout the Indian Ocean region. The Kenya Wildlife
Service recorded 60 whale sharks, with a conservative population estimate of
219, whilst carrying out an aerial survey of marine mammals and turtles in
November 1994 (N. Muthiga, pers. comm.). The sharks were distributed
all along the Kenyan coastline but with a higher frequency of sightings in
northern waters.
Whale sharks are often associated with schools of pelagic fish that are
probably feeding on the same prey organisms. There are numerous references in
the literature to sightings of whale sharks in association with several tuna and
trevally species, and with bonito, mackerel and schools of small bait fish such as
sardines and anchovies. These associations could have foraging advantages for
the whale sharks. Wolfson (1987) reported the sighting of a whale shark
swimming amongst a school of more than 500 hammerhead sharks Sphyrna
lewini (Grith & Smith) oBaja California, and Arnbom & Papastavrou
(1988) observed tiger sharks Galeocerdo cuvieri (Peron & Lesueur) swimming
close to whale sharks in the Galapagos Islands. Groups of whale sharks
have been observed swimming with manta rays Manta birostris (Walbaum) o
Baja California (Wolfson, 1987) and othe Zuytdorp Clis, north of Kalbarri
in Western Australia (P. Weiland, pers. comm.). Other common associates
include several species of remora Remora spp., and the pilot fish Naucrates
ductor (L.). At Ningaloo Reef, Taylor (1994) reported that whale sharks are
often seen accompanied by juvenile golden trevally Gnathanodon speciosus
In the past, the whale shark has been of little interest to man, as it poses no
threat nor is it widely exploited for human consumption or for other products.
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Consequently, there has been virtually no sustained scientific research on this
species and it has been the target of only limited commercial fisheries in the past.
The flesh is soft and bland and has a very high water content, with levels of 68%
(Satyanarayana Rao, 1986) and 75% (A. Giord, pers. comm.) being reported.
Whale shark meat is sought after in Taiwan (Joung et al., 1996), where it is
described as being like tofu and fetches a high price, and currently two
Taiwanese whale shark fisheries are believed to take approximately 100 sharks
annually (Uchida, 1984; J. Stevens, pers. comm.).
Sudhakara Rao (1986), recorded the landing of 40 whale sharks over a
4-day period in 1982, from a harpoon fishery othe Veraval coast in
India. After removal of the liver the carcasses were discarded as there was no
local demand for the flesh. In the past, harpoon fisheries have been reported
from India (Sudhakara Rao, 1986), Pakistan (Anon., 1955;Silas, 1986),
Indonesia (Muller, 1995) and Iraq (Mahdi, 1971). A seasonal (April to May)
fishery exists in the Philippines, where 90 sharks were taken during the
1996 season (Trono, 1996). The anti-tumorogenic properties of whale shark
liver oil have been investigated in China (Zhang et al., 1988). There may
be a developing market for whale shark fins, with reports that some may
recently have been sold in Hong Kong (Smith, 1996). In the Maldives, the
limited fishery for liver oil (Anderson & Ahmed, 1993) has ceased in recent
years, and in 1995 the Ministry of Fisheries and Agriculture introduced
legislation banning all fishing for whale sharks (C. Anderson, pers. comm.).
This protection was introduced because of the low monetary value of the
fishery, the possible serious impact that the fishery may have been having on
whale shark stocks, and the likely benefits to the tuna fishery and tourist
Occasionally whale sharks are taken accidentally in gill and purse seine net
fisheries othe coast of India (Silas, 1986;Devadoss et al., 1990;Seshagiri Rao,
1992). Usually these specimens are discarded, as neither the flesh nor other
products are in high demand. Occasionally some of the flesh is eaten either fresh
or salted and dried, and the liver oil is utilized for water-proofing wooden fishing
boats and other appliances, for the manufacture of shoe polish (Satyanarayana
Rao, 1986) and as a treatment for some skin diseases (Karbhari & Josekutty,
1986). The processing of whale shark fins and fin rays has been reported in India
(Ramachandran & Sankar, 1990).
Often the species is used as a fish ‘ aggregator ’ or indicator of waters rich in
plankton and plankton-feeding fish that will, in turn, attract more valuable
species such as tuna. In the Gulf of Guinea and elsewhere tuna purse seiners seek
out whale sharks on the surface and set nets on them to catch associated fish
species (Stretta & Slepoukha, 1983). However, potential damage to fishing gear
from entangled whale sharks causes them to be avoided in other areas. Several
whale sharks have been kept in captivity in aquaria in Japan (Roth, 1986;
Uchida et al., 1990), but few details are available. There have been a few cases
reported of whale sharks inadvertently ramming boats (Smith, 1967), but
generally the sharks are more at risk from being struck accidentally by vessels
whilst basking or feeding on the surface. There are numerous reports, from the
first half of this century, of collisions between steam ships and whale sharks
(Anon., 1962;Gudger, 1938a,b,1940).
1228 ..
Since 1982, whale shark enthusiast G. Taylor has been studying and photo-
graphing whale sharks at the Ningaloo Reef (Fig. 1). He was the first person to
document the seasonal aggregation of whale sharks in the area, initially through
the collection of informal sightings records from boats. Taylor conducted
preliminary population surveys, and investigated the relationships between mass
coral spawning, the Leeuwin Current, and whale shark occurrences and move-
ment patterns. With funding from the Australian National Parks and Wildlife
Service (ANPWS) he carried out aerial surveys between 1989 and 1992, which
suggested that whale sharks congregate along the reef front during the autumn
months, shortly after synchronous mass coral spawning episodes (Taylor, 1996).
Further work involved limited plankton sampling to try and identify prey
species, observations of feeding behaviour, and initial satellite tracking studies
(Taylor, 1994).
In order to gather more information about population size and sex ratio and
to investigate whether the same animals revisit Ningaloo on a seasonal basis,
Taylor conducted a limited tagging study using conventional game fish tags.
This study, with individuals being re-identified by the location of the tag rather
than its number, was initiated in 1992 and a total of 25 individuals have been
tagged (Taylor, 1994). His estimates of local population size, based upon the
frequency of re-sightings and an estimated tag shedding percentage of 40–50%,
are between 200 and 300 individuals (G. Taylor, pers. comm.). Taylor also
recorded the sex of whale sharks encountered on the reef and investigated the
feasibility of identifying individuals through scars, deformities and the pattern of
spots and stripes behind the gill slits. He has used gross scarring to verify the
long-term stability of the lateral markings and has suggested that these distinc-
tive patterns can be used as a repeatable method of identifying individuals
(Taylor, 1994). He has compiled a photo-identification library, which contains
photographic records for 162 individuals, and it is apparent from these data
that some individual sharks are re-sighted at Ningaloo in successive seasons
(G. Taylor, pers. comm.).
In addition, Taylor has undertaken a long-term study to measure the dorsal
fins of whale sharks at Ningaloo. Repeated re-measurement of the height of the
first dorsal fin of individual sharks, over a period of 10 years or more, may
provide an indication of the growth rate of the fin. If there is a relationship
between this parameter and overall size of the shark, Taylor believes it may be
possible to estimate age of the sharks and to determine when they reach sexual
In 1994, researchers from the Commonwealth Scientific and Industrial
Research Organisation (CSIRO) Division of Fisheries tracked two tagged whale
sharks successfully (one for a period of 26 h) using acoustic telemetry, and
attached archival tags to six individuals, one of which was recovered after 24 h.
The archival or ‘ smart ’ tags can collect data on the date, time, the shark’s
position and swimming depth, and the water temperature for up to 5 years,
giving a record of the shark’s long-term movements once the tag is retrieved
(J. Stevens, pers. comm.). Detailed information on the short-term movements of
the sharks was obtained from the acoustic telemetry tracking and from the single
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archival tag retrieved (Stevens, 1994). Swimming speed ranged from approxi-
mately 0·1 to 1·5 m s
, with the fastest speeds being recorded during the night.
Data from both the acoustic telemetry and the archival tag revealed that the
tagged sharks made numerous dives throughout the 24-h period, to maximum
depths of 90 m, often to within a few metres of the bottom. Diving behaviour of
these individuals did not appear to be linked to the location of the thermocline,
with the acoustically tracked sharks spending the majority of the time above the
thermocline, and the shark tagged with an archival tag spending the day above
the thermocline and most of the night either within or below it. Some of the
tracks revealed that the sharks were circling on the surface oa reef passage,
during ebb tides when water was flowing out of the lagoon, possibly to take
advantage of aggregations of prey associated with mats of algae (J. Stevens, pers.
The accessibility of the seasonal aggregation of whale sharks at the Ningaloo
Reef provides an excellent opportunity for researchers to undertake studies of
this rarely encountered and poorly understood shark. Initial research eorts
lacked clearly defined objectives and were often hampered by limited scientific
knowledge and resources. Dedicated and sustained research of whale sharks
in the Ningaloo Marine Park should not only seek to improve the knowledge of
the whale shark’s biology and ecology, but also to provide information to
managers in order to minimize possible detrimental impacts of tourism pressure.
As suggested by Wolfson (1986), researchers studying whale sharks should
move beyond the purely descriptive natural history approach and design and
implement sustained programmes of investigation, using the most advanced
equipment and techniques that are amenable to statistical treatment ’.
In developing a scientifically objective research and monitoring programme,
there are a number of factors to be considered, including ethical, technical and
logistical issues. The large size, free-swimming epipelagic nature, and sporadic
appearance of whale sharks makes study of these animals intrinsically dicult
and creates numerous technical and methodological problems. As has been seen
with cetacean research, the time required and complexity of programmes
examining any large marine animal have to be considered at the design stage.
There are problems in initiating further research in an area where a whale shark
watching industry is already in place. Attempts to investigate the population size
and structure may suer from sample size and range problems. Population
studies require large sample sizes and this is a major problem when working with
rarely encountered species, especially if individuals cannot be captured or
restrained. Accurate morphometric data and samples for age and growth rate
determination (such as vertebral centra) can usually be obtained only from
restrained or dead animals.
A key factor in sustainable management of whale shark–human interactions is
a clear understanding of the population dynamics of the animal. Until both
seasonal and interannual variability in abundance and distribution are known, it
will be dicult to identify any long-term impacts. Therefore, monitoring studies
have to establish an independent and repeatable series of population counts.
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Reliable estimates of whale shark population size at Ningaloo are dicult to
obtain. In the past there has been no standardized collection of data. If
segregation by age occurs, which appears to be the case from anecdotal evidence,
sightings in the area are not a random sample of the population. Habituation to
boats may occur, which also negates random sampling. A sucient sample size
is required also. At present it is impossible to fix the spatial boundaries of the
population, as there is no indication where the sharks may migrate. A long-term
aerial survey programme to monitor interannual variability in the whale shark
population is a high priority for future management of human–whale shark
interaction in the Ningaloo Marine Park.
My thanks to C. Simpson, J. Burt and J. Stevens for reviewing the manuscript, to R.
Laurie for supplying the map of the Ningaloo Marine Park and to all those who provided
information for this review, including C. Anderson, G. Cli,A.Giord, D. Goorah,
S. Menzies, N. Muthiga, N. Nannup, R. Salm, G. Taylor and P. Wieland.
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... The whale shark Rhincodon typus (Orectolobiformes: Rhincodontidae) is the largest elasmobranches found in the tropical and temperate seas worldwide including coastal, neritic and pelagic habitat (Colman 1997;Couturier et al. 2012). Despite their wide range of distribution, very little is known of their pelagic distribution. ...
... On both occasions, R. typus has sighted approximately at 5-10m depth below the sea surface. Whale sharks distribution and biology are very meager in oceanic waters whale sharks are susceptible to worldwide exploitation ranging from incidental by-catch in fisheries to direct capture in some regions (Colman 1997;Hanfee 2001). In the present observations provide important information on the occurrence and distribution of this species in the oceanic waters of India. ...
... In the present observations provide important information on the occurrence and distribution of this species in the oceanic waters of India. The distribution and abundance of whale sharks is known to be influenced by oceanographic factors such as sea surface temperature, salinity, current and primary productivity (Colman 1997;Hoffmayer et al. 2005). Krishnapatnam Coast has dynamic oceanographic features and high productivity of this area. ...
The present study was based on sighting of whale shark, Rhincodon typus Smith, 1828 on 13th and 14th October 2016 incidentally in oceanic waters of Bay of Bengal at a depth of 3340 and 3270 m. These two sightings were thus confirmed by photographs. Whale sharks were reported in the coastal and near shore regions of India by many of the researchers and this is the first observation of R. typus in oceanic waters of east coast of India.
... Las migraciones parecen estar relacionadas con cambios en la temperatura de las masas de agua y con la búsqueda de alimento, es por ello que se dirigen hacia zonas con alta productividad primaria, donde la especie se agrega por periodos predecibles y largos de tiempo (Compagno 1984;Clark y Nelson 1997;Colman 1997;Heyman et al. 2001;Wilson et al. 2001;Ketchum et al. 2012). ...
... Nacen pequeños (menos de un metro) aumentando las posibilidades de ser depredados. Los primeros registros de neonatos fueron en el estómago de marlín (Colman 1997) y Tiburón azul (Kukuyev 1996). Además, existe un registro de orcas comiendo un Tiburón ballena de ocho metros (O´Sullivan 2000). ...
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La presente obra es resultado de la participación de expertos provenientes del sector gubernamental, académico y privado, con un total de 36 autores que compilaron información contenida en investigaciones científicas y material bibliográfico (tesis de posgrado, libros y artículos científicos). Sin duda, el contenido facilitará significativamente la labor de la Autoridad Científica cites de México, pues aporta elementos para analizar y emitir DENP de manera oportuna, además de apoyar a las autoridades Administrativa y de Aplicación de la Ley de México para su efectiva implementación de la cites, cuyo tratado internacional es uno de los más efectivos para la gestión de nuestro patrimonio natural, del cual dependen la subsistencia y modos de vida en el largo plazo de las comunidades pesqueras.
... Las migraciones parecen estar relacionadas con cambios en la temperatura de las masas de agua y con la búsqueda de alimento, es por ello que se dirigen hacia zonas con alta productividad primaria, donde la especie se agrega por periodos predecibles y largos de tiempo (Compagno 1984;Clark y Nelson 1997;Colman 1997;Heyman et al. 2001;Wilson et al. 2001;Ketchum et al. 2012). ...
... Nacen pequeños (menos de un metro) aumentando las posibilidades de ser depredados. Los primeros registros de neonatos fueron en el estómago de marlín (Colman 1997) y Tiburón azul (Kukuyev 1996). Además, existe un registro de orcas comiendo un Tiburón ballena de ocho metros (O´Sullivan 2000). ...
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El Tiburón sedoso, Carcharhinus falciformis, tiene una distribución circunglobal, se localiza tanto en el Atlántico Occidental y Pacífico Oriental. En México, C. falciformis se encuentra en el Pacífico Mexicano, incluyendo el Golfo de California, así como en el litoral del Golfo de México y Mar Caribe. Los tiburones sedosos habitan generalmente plataformas continentales e insulares, pendientes e incluso se han registrado ocasionalmente en aguas someras (18 m) hasta profundidades de 550 m. La edad máxima estimada para C. falciformis varía considerablemente para las distintas poblaciones, desde ocho a 11 años hasta más de 30 años. Los parámetros de crecimiento son también altamente variables, con longitudes asintóticas que van de 240 cm a 340 cm de LT (longitud total). El Tiburón sedoso presenta una estrategia de reproducción vivípara placentaria, con un periodo de gestación de 11 a 12 meses y una fecundidad de una a 25 crías con tallas entre 50 y 83 cm (LT). En el 2014 C. falciformis fue enlistada en el Apéndice II de la Convención sobre la Conservación de las Especies Migratorias de Animales Silvestres (CMS) y en 2017 se incluyó en el Apéndice II de la Convención sobre el Comercio Internacional de Especies Amenazadas de Fauna y Flora Silvestres (CITES). En México, la falta de información de captura y esfuerzo pesquero específico ha complicado la evaluación del estado de las poblaciones de tiburones, en donde C. falciformis no es la excepción. Por ello, es necesario hacer registros específicos de las capturas, esfuerzo pesquero, datos biológicos-pesqueros, con el fin de realizar análisis demográficos y proyecciones del efecto de la pesca en el stock e identificar áreas de importancia para su protección.
... Maturity in females could not be visually assessed, but females ≥9.0 m were assumed to be mature, falling in line with previous studies of whale shark demographics (Colman, 1997;Meekan et al., 2020). ...
The world's largest fish, the whale shark (Rhincodon typus), is a circum‐tropically distributed and globally endangered species, that is widely studied at predictable aggregation sites. The main Hawaiian Islands are not known to have large aggregations of whale sharks; however, they have been anecdotally reported here with some regularity. To date, little is known about this charismatic species in Hawaiian waters. This study is the first effort to examine whale shark demographics and movements in a previously unstudied region of the world. Here, citizen science was used to investigate the abundance, seasonality (if any) and occurrence of whale sharks in the waters around the main Hawaiian Islands. A total of 309 individual whale sharks were identified from sightings between 1991–2020 by their unique spot patterns, most of which (74%) were reported between 2018 and 2020. Estimated whale shark total length ranged from 2–12 m (mean 6.1 m), with both juvenile and mature males and females represented in the dataset. The best‐fit Lagged Identification Rate (LIR) model for these data suggests that individuals sighted in Hawaiian waters are transient in nature, supporting the empirical data (88% of sharks sighted only once). Although using citizen science data can have inherent biases, these results present a conservative first assessment of the demographics of whale sharks in Hawaiian waters. For an Endangered species assessed as Largely Depleted by the IUCN Green Status of Species, highlighting Hawai’i as an important habitat for the whale shark – mainly as a migratory corridor or navigational waypoint – is a crucial step in understanding their ecology in the Pacific in order to develop effective management plans.
... While even the larger R. typus may occasionally be prey to large marine predators, particularly Orcinus orca (Killer Whale), and possibly large sharks (Carcharhinidae and Lamnidae) (Rowat and Brooks 2012), it is while individuals are still relatively small (<2 m) that they are likely to be most vulnerable. Small individuals have been found in the stomachs of Makaira mazara (Indo-Pacific Blue Marlin) and Prionace glauca (Blue Shark) (Colman 1997). The uniquely large litter size of R. typus among live-bearing shark species, which can be ~300 pups ( Joung et al. 1996), suggests an evolutionary strategy to counter a high pup mortality rate. ...
Full-text available
This chapter in the New Natural History of Madagascar (Goodman ed. 2022, Princeton University Press) provides the most comprehensive account to date of scientific knowledge of the natural history of Madagascar's marine and coastal ecosystems and their biodiversity, including marine megafauna (including cetaceans, dugong, sea turtles, sharks, rays, sawfishes and coelacanth (Latimeria chalumnae).
... The stranded specimen was larger than some of the specimens tagged by Robinson et al. (2017) in Qatari waters of the Arabian Gulf, but the two specimens sighted in the Shatt al-Arab Estuary were much smaller than those studied by Robinson et al. (2016Robinson et al. ( , 2017. Babu et al. Colman (1997) stated that the record by Mahdi (1971) from the marine waters of Iraq was based on harpoon fisheries, but it is clear from the original paper that no specimen was captured and only a sighting was given. Al-Shammary (2014) published two images of R. typus in his record, but one of them is clearly taken from a previous publication. ...
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This study is the first record of a whale shark Rhincodon typus in Iraqi waters and the second record from the Arabian (Persian) Gulf portion of the northwest Indian Ocean. The stranding and sighting events were documented by photos of three individuals: the stranded specimen was 6.5 m total length (L T) and the free-swimming individuals were c. 1-2 and 2-3 m L T respectively . An appeal is made for Iraqi policy makers to participate in regional and international organizations for the conservation of this endangered species.
... Hiu paus tergolong jenis ikan perenang cepat dan memiliki karaktersitik biologi yang berbeda dengan kelompok hiu lainnya (Venegas et al., 2011) dengan wilayah penyebaran yang luas (Sequeira et al., 2012), namun pengetahuan tentang biologi dan ekologi hiu paus masih sangat terbatas (Syah et al., 2018). Sebaran hiu paus terdapat di perairan tropis dan sub tropis yang hangat (suhu berkisar 18-30 ᵒC) di antara 30 ᵒ Utara dan 30ᵒ Selatan (Colman, 1997;Tania & Noor, 2014). Di Indonesia, kemunculan hiu paus dapat ditemukan pada beberapa perairan dengan periode waktu tertentu, seperti perairan Pangandaran yang muncul pada bulan Agustus-September dan di perairan NTT pada bulan Agustus-November (Stevens et al., 2007;Kamal et al., 2016). ...
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Hiu paus (Rhincodon typus) adalah ikan terbesar di dunia yang sangat berpotensi dalam bidang wisata, namun potensi ancamannya juga tinggi jika tidak dikelola dengan baik. Perairan Kwatisore merupakan salah satu habitat hiu paus di Indonesia yang terlihat sering muncul. Hal tersebut menjadi fenomena yang unik karena hiu paus muncul setiap hari dan sepanjang tahun, sehingga berpeluang untuk pengembangan wisata. Penelitian ini bertujuan untuk mengkaji karakteristik pola makan hiu paus dan agregasinya, terutama total jumlah kemunculan dan jumlah individu di perairan Kwatisore. Pengambilan data dalam penelitian ini berupa data primer dan dilakukan dari bulan Juli sampai September 2020. Metode pengambilan data dilakukan sekali setiap bulan (time series) melalui koleksi untuk mengkaji pola makan antar individu hiu paus dengan pengamatan secara langsung ketika hiu paus naik ke permukaan perairan dan berada di dalam bagan yang berjumlah 5 unit alat tangkap bagan. Hasil penelitian menunjukkan bahwa terjadi 275 kemunculan hiu paus, yang terdiri dari 18 individu. Jumlah kemunculan dan jumlah individu hiu paus sangat tergantung pada hasil tangkapan ikan teri oleh setiap unit alat tangkap bagan. Kemunculan hiu paus berada dalam petuanan hak ulayat laut Kampung Akudiomi. Kemunculan hiu paus dominan terjadi pada pagi hari dan persentase kemunculan 100% berjenis kelamin jantan dengan ukuran panjang total berkisar antara 3 hingga 7,5 m.
... An ecotoxicological investigation using skin biopsies was performed on the whale shark (R. typus) in the Gulf of California (Mexico) (Fossi et al., 2017). This species has a circumequatorial distribution in all tropical and warm temperate seas (Compagno, 1984;Colman, 1997) and it is the largest living fish, which grows up to 20 m. In 2000 whale sharks were declared as vulnerable according to the IUCN Red List (Norman, 2000) and in 2016 their conservation status was assessed as endangered (Pierce and Norman, 2016). ...
Due to the recognized negative impact for marine biodiversity, plastic pollution in seas and oceans is nowadays a global problem. Sharks and rays are apex predators, which are also vulnerable to plastic pollution due to threats related with entanglement, ingestion, and habitat degradation, but, paradoxically, the science community deserved them less attention. Entanglement with ghost nets and other plastic materials such as ropes, straps, bags is common, and many evidences of plastic ingestion in shark are also reported. Large sized planktivorous sharks and mobulid rays are seriously exposed to microplastic pollution due to their feeding mode. Despite the occurrence of plastic pollution, elasmobranchs seem to be much more threatened by the effects of the overexploitation through targeted fisheries and by-catch than other stressors. Best practices and environmental laws together with a wide popular participation are necessary to overcome these problems.
Abstract Filter feeding elasmobranchs may be considered as biological indicators of marine pollution, despite most of these species are under some degree of extinction risk. Among threats to this taxonomic group, marine pollution might represent an additional concern for their survival. In this review, a comprehensive systematic search of scientific literature on pollutants in filter feeding elasmobranchs was conducted to evaluate the bioaccumulation patterns, and risk for human consumers. We found that, despite an increasing trend in the number of published studies, the geographical coverage is still very limited and most of the studies focused solely on trace elements (70.8%). Among sharks, Rhincodon typus was the most represented species (66.7%), while Mobula mobular the most studied ray species (41.7%). Comparing the levels of pollutants in filter feeders between ocean basins, this review highlighted that Hg, As and Cd levels are mostly higher in those areas affected by both strong natural and anthropogenic source of emissions, such as the Indian Ocean. With regards to OCs, ΣPCB levels in muscle of C. maximus were between 4.3 and 50.5 μg kg−1 ww, highlighting a persistent contamination of PCB in the Mediterranean Sea. Some species exceeded the maximum allowable limits for foodstuff consumption for As, Cd and Pb. A total of 77.8% of the analyzed species exceeded the Environmental Quality Standards for Hg, while they were always below the EQSbiota for HCB, PBDEs, PFOS and DDT. Given their feeding mechanism that continuously samples the marine environment, further investigations are urgently needed to determine not only the extent of contaminant exposure in different hotspot locations but also the risks posed to the elasmobranch health.
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Whale sharks (Rhincodon typus) are found circumglobally in tropical, subtropical, and warm temperate waters, and their known seasonal aggregations and migratory movements are influenced by factors such as ocean currents, thermobiological systems, and patterns of productivity. Several locations in the eastern tropical Pacific Ocean are known habitats for R. typus; Although it has long been known that whale sharks aggregate along the Panama coast, little is known in relation to their movement patterns, behavior, and habitat use. In this study, we investigated the movements and behaviors of R. typus tagged in Panama in relation to oceanographic variables and examined the overlap of foraging habitat and migratory routes with marine protected areas (MPAs), industrial fishing areas, and marine traffic. Satellite tracks from 30 R. typus tagged in the coastal waters of Panama were examined, including nine tags suspicious of earlier detachment. A hidden Markov model was then used to identify different behavioral states (foraging and migrating) and their relationships with environmental variables (sea surface temperature, primary productivity, chlorophyll-a concentrations, and eddy location/speed) Tracks were also superimposed on maps of MPAs, industrial fishing areas, and regional marine vessel traffic to identify the degree of overlap. Rhincodon typus foraged mainly within the Panamanian exclusive economic zone but also moved north and south along the coast and out to the open ocean. Significant differences in environmental conditions were found between sites in which foraging and migrating behaviors were recorded. Higher productivity and chlorophyl concentration were associated with foraging behavior, while higher eddy speeds were observed when sharks migrated. Rhincodon typus used MPAs; however, there was a high degree of overlap between their habitat and areas of industrial fishing and marine vessel traffic. Our results highlight the use of the coastal waters of Panama, oceanic seamounts, and ridges, MPAs and industrial fishing areas by R. typus for foraging and migration. Additionally, our findings highlight the importance of satellite tracking studies for understanding the behavior and habitat use of highly mobile migratory species, such as R. typus.
Mass spawning occurred mainly around the third quarter of the moon on neap, nocturnal, ebb tides. Some 102 species of scleractinian corals from Western Australian Reefs are known to spawn during the austral autumn. A further 44 species were found to contain ripe gonads during the same period and are presumed to participate in the annual coral mass spawning on Western Australian reefs. These records represent 88% of the coral species studied so far or about 46% of the coral species currently described from Western Australia. In Western Australia, coral mass spawning coincides approximately with the annual intensification of the Leeuwin Current, a warm poleward current of tropical origin that flows along the coastline of Western Australia during the austral autumn and winter. This current provides a mechanism for the southward dispersal of planulae and raises the possibility of a unidirectional gene flow between regionally separate coral reefs in Western Australia. Comparisons with the spring coral mass spawnings on the Great Barrier Reef indicate that, apart from the seasonal difference in the timing of spawning, many similarities exist suggesting that the same phenomenon is occurring on both sides of Australia. The different seasonal timing of coral mass spawning on the E and W coasts of Australia is probably the result of an endogenous rhythm reflecting the breeding patterns of ancestral corals as a consequence of selective dispersal of larvae from equatorial regions. -from Author