Stenella coeruleoalba (Meyen,
English: Striped dolphin, blue-white dolphin
German: Blau-Weißer Delphin
Spanish: Delfín listado
French: Dauphin bleu et blanc, dauphin rayé
Stenella coeruleoalba © Würtz-Artescienza (see "links")
The species name "coeruleoalba" refers to the
pattern of blue/dark-gray and white stripes and blazes along the
lateral and dorsal sides of the body. The dorsal cape is muted blue
or blueish-grey, usually invaded by a white to light grey spinal
blaze. The sides are darker than the belly. Striped dolphins have
a long beak, well demarcated from the melon and falcate dorsal fin.
In the field, they are most likely confused with common dolphins
delphis) and other similar-sized species but can be distinguished
by their robust body and coloration. The largest recorded specimen
was 2.56 m long and the maximum weight recorded was 156 kg. Mean
body length in the western Pacific is 2.4 m for males and 2.2 m
for females (Archer, 2009).
Striped dolphins show only moderate geographical variation in skeletal
morphometrics and little if any geographical variation in pigmentation
pattern. However, several authors found slight but significant differences
in body size between local populations in the eastern North Atlantic,
the northwestern Mediterranean, and the southwestern Mediterranean
(Rice, 1998). MtDNA and microsattelite differentiation suggests
that NE-Atlantic striped dolphins form a separate population from
the Mediterranean population (Garcia-Martinez et al. 1999; Bourret
et al. 2007).
The striped dolphin is distributed world-wide in tropical and temperate
waters. It ranges north in the Atlantic to Newfoundland, northern
Scotland, and Denmark; in the Mediterranean Sea; and in the Pacific
to the Sea of Japan, Hokkaido, about 40°N across the western
and central Pacific, and British Columbia (Canada). The southern
limit of its range is in Argentina, Cape Province of South Africa,
Southern Australia, New Zealand, and Peru (Archer and Perrin, 1999;
Hammond et al. 2008).
Distribution of S. coeruleoalba (Hammond et al.
2008; © IUCN; enlarge
warm temperate, subtropical, and tropical waters around the world.
Although Perrin et al (1994) stated that it is not
a common inhabitant of cold boreal waters as previously claimed,
there are coldwater records, e.g. from Greenland and the Faroe Islands,
and Syvertsen et al. (1999) and Isaksen and Syvertsen (2002) reported
sightings/strandings from the Norwegian and Swedish coasts. Vagrants
have even been recorded from Komandorskiye Ostrova (Rice, 1998).
3. Population size
Würsig et al. (1998) assessed cetacean responses to survey
ships and aircraft and found that S. coeruleoalba moved to
avoid the ships in 33% of sightings. This indicates that density
estimates for this species may tend to be biased downwards.
In the western Atlantic Ocean, the most recent estimate of abundance
for striped dolphins in oceanic waters of the Northern Gulf of Mexico
(US EEZ) pooled from 2003 to 2004, is 3,325 animals (CV=0.48) (Mullin
2007). This value is not statistically different from the 1996-2001
estimate of 6,505 (Warring et al. 2009).
In the eastern Atlantic Ocean, the Bay of Biscay population size
was estimated at 74,000 animals in 1993 (Goujon, 1996) and more
recently at 56 500 (95% CI 29 100-90 400) in 2002 (Certain et al.
In the central Mediterranean, striped dolphins were the most abundant
species (43.5%) observed during surveys (Perrin et al. 1994, Reyes,
1991 and refs. therein). In the waters of the central Spanish Mediterranean
coast, 2001 - 2003 estimates yield a mean abundance of 15,778 dolphins
(95% CI = 10,940-22,756). This density is comparable to that obtained
in the International Ligurian Sea Cetacean Sanctuary (Gomez et al.
2006). In the southern Tyrrhenian Sea around the Aeolian archipelago
the 2003 estimate was 4,030 individuals (CV=0.30) (Fortuna et al.
In the Pacific Ocean, the preliminary 2003 estimate of abundance
for the 2001-2005 striped dolphins is 1,470,854 (CV=0.15) animals
(Gerrodette et al. 2005). Geometric mean abundance estimate for
California, Oregon and Washington waters based on the 2001 and 2005
ship surveys is 17,925 (CV=0.37) striped dolphins (Forney 2007).
A 2002 shipboard line-transect survey of the entire Hawaiian Islands
EEZ resulted in an abundance estimate of 10,385 (CV=0.48) striped
dolphins (Barlow 2003). This is currently the best available abundance
estimate for this stock.
In the western North Pacific, most recent estimates are dated (Hammond
et al. 2008) and stem from the 1980's: 570,000 (CV = 0,19) striped
dolphins occurred there (Miyashita, 1993). Two areas of concentration
were identified, comprising about 52,682 animals (CV = 95%) between
20° and 30°N and around 497,725 (CV = 18%) animals between
30° and 40°N. Relatively few striped dolphins (about 19,631;
CV = 70%) were present in the nearshore waters off Japan. However,
there is clear evidence that there has been a decline since the
1950's as a result of stock depletion by over-hunting (Kasuya 1999).
They are uncommon in the Sea of Japan, East China Sea and Ryukyuan
waters (Reyes, 1991).
There is very little data from other areas of distributional range:
Ballance and Pitman (1998) found that S. coeruleoalba was the second-most
abundant species sighted in the western tropical Indian Ocean (14%
of all cetaceans, compared to 33% for the eastern tropical Pacific
and 10% for the Gulf of Mexico). Striped dolphins are also reported
from the coast of Angola (Weir, 2007).
4. Biology and Behaviour
Habitat: Striped dolphins are pelagic animals. In the Pacific
waters off Japan, they are found in three geographical aggregations
in the summer, between 20 and 42 °N. Occurrence is seasonal
in the northern part of the range (Toshio 1999). In the eastern
Pacific, they prefer areas with large seasonal changes in surface
temperature and thermocline depth and with seasonal upwelling (Reyes,
In western North Atlantic waters, striped dolphins seem to be confined
to the Gulf Stream or the waters off the continental slope (Davis
et al. 1998). Along the Mid-Atlantic Ridge from Iceland to the Azores
they inhabit warmer (12-22 °C) and more-saline (34.8-36.7 psu)
waters in the south (Doksaeter et al. 2008).In the eastern North
Atlantic, as well as off South Africa they are found in deep water
(greater than 1,000m) past the continental slope (Perrin et al.
1994 and refs. therein). In the Strait of Gibraltar and in the Mediterranean
they prefer waters of 600 m or more depth (Hashmi, 1990; Reyes,
1991; Bourreau and Gannier, 2003).
Schooling: Schools are of varying size and composition.
Of 45 schools examined from off the coast of Japan, most (86%) contained
fewer than 500 individuals. The mean school size was 101 animals.
Schools moving south with the retreating front of the Kuroshio Current
are larger than those moving north earlier in the year. Schools
in the eastern North Atlantic more commonly have 10-30 individuals
and rarely reach the hundreds. In the western Pacific, three major
types of schools are recognised: juvenile, adult and mixed, the
latter being divided into breeding and non-breeding schools. Juvenile
schools may migrate closer to the coast than adult and mixed schools.
Calves remain in adult schools until 1 or 2 years after weaning
and then leave to join juvenile schools (Perrin et al. 1994 and
Food: : Feeding depth may extend to below 200m and down
to 700m (Archer, 2009); 75-80% of the prey in the Japanese and South
African material had organs of luminescence. Individual fish in
the stomachs of the animals captured off Japan ranged in length
from 60 to 300mm (Perrin et al. 1994 and refs. therein; Santos et
al. 2001a, 2001b). Myctophid fish predominate in specimens from
Japan and South Africa whereas in dolphins stranded on the Mediterranean
coasts of France, Spain, and Italy, cephalopods dominate in the
stomach contents. Blanco et al. (1995) found that the cephalopods
Albraliopsis pfefferi, Onychoteuthis banksii, Todarodes
sagittatus and Brachioteuthis riisei were dominant in
stomach samples from the western Mediterranean. In the eastern Mediterranean
Sea off the Turkish coast, the cephalopod Abralia veranyi
was the most common prey (51.2% of all the beaks found in this species),
followed by Onychoteuthis banksii and Heteroteuthis dispar.
The stomachs also contained remains of fish and shrimp (Özturk
et al. 2007).
In the oceanic waters of the Northeast Atlantic, the diet was found
to be primarily composed of fish (39% by mass) and cephalopods (56%)
and less of crustaceans (5%). The most significant fish family identified
was the lanternfish (24%). The oceanic squid Teuthowenia megalops
and Histioteuthis spp. were the most significant. The pelagic
shrimp Sergastes arcticus and Pasiphaea multidentata
were the most prevalent crustaceans. Prey sizes ranging from 30
to 170 mm accounted for 80% of the prey items, while 80% of the
reconstituted biomass consisted of prey measuring between 60 and
270 mm. Prey composition and size-range differed slightly with sex
and age or body size of the dolphins. The state of digestion of
food remains suggested that predation took place at dusk or during
the early hours of the night (Ringelstein et al. 2006). In the Bay
of Biscay striped dolphins are able to shift from vertically migrating
meso-pelagic prey to neritic or coastal prey types (Spitz et al.
2006). The diet of striped dolphins also varies according to food
availability both in terms of quantity and composition, reflecting
changes in the relative abundance of fish species (Spiz et al. 2003).
While in some regions (e.g. portions of the US east coast) striped
dolphins are encountered in all seasons, they elsewhere appear to
be associated with the fronts of warm oceanic currents that move
seasonally and produce sporadic warm-water intrusions and meanders.
In Japanese waters, the species is associated with the northern
boundary of the warm Kuroshio Current, which extends up to 46°N
in the summer and retreats to 33°N in the winter. It appears
earlier in the season than S.
attenuata, consistent with the hypothesis that the latter
is the more tropical (Perrin et al. 1994 and refs. therein). Striped
dolphins approach the coast in September and October and move southward
along the coast, apparently dispersing into the East China Sea for
the winter. In April they return along roughly the same route, but
farther offshore. Eventually they leave the coast to summer in the
pelagic North Pacific. Segregation by age is observed (Reyes, 1991).
Seasonal movements may also occur in the Mediterranean. The dolphins
move towards the northern part of the basin as the sea surface temperatures
in the southern part increase. Between the French mainland and Corsica,
data obtained 2001 - 2004 show that relative abundance peaks in
May and September, while a consistent minimum is obtained from December
to April (Laran and Drouot-Dulau, 2007).Sighting data also suggest
seasonal movements of this species in the eastern tropical Pacific
(Perrin et al. 1994; Reyes, 1991 and refs. therein).
In the Ligurian Sea of the French Riviera, there are also diurnal
offshore-inshore movements. Night acoustic results show the presence
and intense feeding activity of striped dolphins close to the shelf
break. Day distribution shows a marked preference for the open sea
Direct catch: The largest direct catches have been taken
in Japanese waters, in drive and hand-harpoon fisheries at several
locations. The catches were voluntarily reduced beginning in 1981
and have since varied between 358 (in 1987) and 4,883 (1981), averaging
2,830 during the period 1981-89. Between 1989 and1993, the average
annual catch was 1,028.
Kasuya (1999) reported that Japanese multispecies dolphin fisheries
now receive an annual quota of 725. Ten years later, striped dolphins
continue to be caught in Japan. In the Wakayama prefecture, 86 were
landed by hand-harpoon and 384 by driving fisheries in 2007. The
quota for 2007/2008 was set at 685 dolphins (Iwasaki, 2008). Fragmented
information on morphology, life history, pollutant levels and genetics
suggests that the striped dolphins taken by Japanese fisheries are
from more than one population, with varying proportions among fisheries
and perhaps over time (Kasuya, 1999).
Striped dolphins are also taken in the drive fishery at Malaita
in the Solomon Islands and in the harpoon fishery for small cetaceans
at St Vincent. Other such small indigenous fisheries may exist elsewhere.
Small numbers were taken by French and Spanish fishermen for human
consumption in the Mediterranean (Perrin et al. 1994 and refs. therein;
Jefferson et al. 1993).
In the Northeast Atlantic, striped and common dolphins were harpooned
to supply food for consumption on board or to scare them away from
tuna trolling lines. It is difficult to ascertain the number of
dolphins taken in this way, but it has been estimated in the thousands
Incidental catch: Incidental catches are known to occur in
gillnets in the north-eastern Indian Ocean, in tuna purse seines
in the eastern tropical Pacific, in fisheries in the northeastern
Atlantic, in drift nets, purse seines and other gear in the Mediterranean,
in various gear off the coast of Japan, and in drift gillnets in
the North Pacific, and probably occur in similar fisheries in tropical
and warm-temperate waters around the world. Although rare, striped
dolphins have also been caught in shark nets in Natal and South
Africa (Perrin et al. 1994 and refs. therein).
Recently, the Inter-American Tropical Tuna Commission estimated
that 6 striped dolphins were caught as by-catch in the 2007 purse-seine
tuna fishery (IATTC, 2009). There has been no reported fishing-related
mortality during 1998-2006 in the US waters of the Northern Gulf
of Mexico or off the northeast U.S. coast (Warring et al. 2009).
Similarly, no striped dolphins were observed killed in the most
recent five-year period in eastern Pacific US waters (Carretta et
Elsewhere, the news are less good. Despite a UN moratorium on the
use of drift-nets in the high seas and a ban in the Mediterranean
by all European Union countries, some fisheries continue to operate
illegally. In 2002 and 2003 line-transect surveys conducted in the
southern Tyrrhenian Sea around the Aeolian archipelago estimated
the striped dolphin by-catch as 36 by-caught animals over a period
of only 12 days. These results are a cause for concern (Fortuna
et al. 2007). According to international official sources, Morocco
harbors the bulk of this fleet in the Mediterranean. Between 2002
and 2003 the driftnet fleet targeting swordfish (Xiphias gladius)
based in Al Hoceima (Alboran Sea) consisted of at least 177 units
with estimated average net length ranging from 6.5 to 7.1 km. This
fleet causes estimated by-catch mortality of 3,110-4,184 common
and striped dolphins per year, and annual take rates exceed 10%
of their population sizes in the Alboran Sea (Tudela et al. 2005).
Antoine et al. (2001) estimated that by-catches in the tuna drift-net
fishery in the Northeast Atlantic were to 90% composed of Delphinus
delphis and S. coeruleoalba. Mean catch rate by trip
in the years 1992-1993 were 4.7 striped dolphins per km of net and
per day. Such rates are similar to those estimated in other driftnet
fisheries. Goujon (1996) estimated the annual additional mortality
linked to the driftnets in the Bay of Biscay albacore tuna fishery
to 1.8% for the striped dolphin (this estimate must be increased
by 30% in order to take into account the whole European albacore
tuna driftnet fishery). The extrapolated decadal scale data from
Irish and other driftnet fleets operating in this area suggest that
during the period 1990-2000, a minimum of 12,635 (10,009-15,261)
striped dolphins were killed as by-catch (Rogan and Mackey, 2007).
Unfortunately, acoustic deterrents developed for harbour porpoise
phocoena) show no effect on striped dolphins (Kastelein
et al. 2006).
In the Southwest Atlantic, off Brazil, by-catch of S. coeroleoalba
was also noted (Zerbini and Kotas, 1998).
Overfishing: The European anchovy is the most heavily exploited
pelagic resource in the Mediterranean, where some other stocks of
pelagic fish are already over-exploited. Since striped dolphins
are reported to eat anchovies and sardines in the area, this could
eventually become either a source of conflict with the commercial
fisheries or a potential threat for dolphin populations (Reyes,
1991). The 1990-1992 epizootic devastated the whole Mediterranean
population; over one thousand corpses were examined in the western
Mediterranean alone, but the toll was probably much higher. The
causative agent of the die-off was a morbillivirus, but the effect
of some pollutants and decreased food availability were suggested
as triggering factors. Depletion of fish and cephalopod resources
is widespread in the Mediterranean and, given that the diet of striped
dolphins includes commercial species, this undoubtedly has a potential
for limiting population numbers (Aguilar, 2000). .
Pollution: Contaminants have been studied more intensively
in this species than in any other cetacean. A long series of papers
has reported the levels, accumulation rates, distribution, relationships
and transfer dynamics of organochlorine compounds and heavy metals
in striped dolphins taken in the Japanese drive fishery or washed
ashore as by-catch in the Mediterranean.
In Japanese samples, increasing trends of PBDEs and HBCDs were observed,
suggesting a growing pollution in Japan and surrounding countries
in recent years (Isobe et al., 2009).
In European waters, decreasing lead concentrations in tissues reflect
the decrease in the production of alkyl lead and the increasing
use of unleaded gasoline (Caurant el al., 2006). Similarly, concentrations
of PCBs, DDT and its metabolites have slowly decreased, although
the decline in PCB has been steeper than that of DDT, suggesting
that the offshore marine environment has not been exposed to significant
releases of these contaminants in recent years (Aguilar and Borell,
2005; Wafo et al., 2005). However, the detected levels reflect the
ubiquity and environmental persistence of these compounds.
Other studies revealed high levels of mercury in striped dolphins
from the Ligurian, Adriatic, and Thyrrenian Seas (Cardellicchio,
2000). Monaci et al. (1998) found that mercury levels were higher
in tissues from animals stranded on the Italian coasts and in skin
biopsies obtained in the Tyrrhenian and Ligurian Seas than in the
respective Spanish samples. This is probably related to Hg pollution
from the natural weathering of cinnabar ores in central Italy. Geographical
differences in trace-element accumulation patterns may also reflect
the existence of two different populations of S. coeruleoalba
in the western Mediterranean.
According to Aguilar (2000) tissue levels of organochlorine compounds,
some heavy metals and selenium are high in Mediterranean samples
and exceed threshold levels above which detrimental effects commonly
appear in mammals. However, apart from the indication that these
levels may have acted as triggering factors in the 1990-1992 epizootic
by depressing the immune system of diseased individuals and potential
lesions in the ovaries, no information on pollutant-related effects
Noise pollution: Observations undertaken during seismic
surveys employing airguns in UK and adjacent waters show a clear
effect on several dolphin species. Small odontocetes showed the
strongest lateral spatial avoidance (extending at least as far as
the limit of visual observation) in response to active airguns (Stone
and Tasker, 2006). In 2005, there was a series of stranding events
throughout Taiwan, involving several species including striped dolphins,
which was linked to navy exercises using sonar (Wang and Yang, 2006).
Range states (Hammond et al., 2008):
Algeria; American Samoa; Anguilla; Antigua and Barbuda; Aruba; Australia;
Bahamas; Bangladesh; Barbados; Belize; Benin; Bermuda; Brazil; British
Indian Ocean Territory; Brunei Darussalam; Cambodia; Cameroon; Canada;
Cape Verde; Cayman Islands; China; Cocos (Keeling) Islands; Colombia;
Comoros; Congo; Congo, The Democratic Republic of the; Cook Islands;
Costa Rica; Côte d'Ivoire; Cuba; Cyprus; Denmark; Djibouti;
Dominica; Dominican Republic; Ecuador; El Salvador; Equatorial Guinea;
Ethiopia; Fiji; France; French Guiana; French Polynesia; Gabon;
Gambia; Germany; Gibraltar; Greece; Greenland; Grenada; Guadeloupe;
Guam; Guatemala; Guinea; Guinea-Bissau; Guyana; Haiti; Honduras;
Hong Kong; India; Indonesia; Iran, Islamic Republic of; Ireland;
Israel; Italy; Jamaica; Japan; Kenya; Kiribati; Kuwait; Liberia;
Madagascar; Malaysia; Maldives; Marshall Islands; Martinique; Mauritania;
Mexico; Micronesia, Federated States of; Monaco; Morocco; Mozambique;
Myanmar; Namibia; Netherlands; Netherlands Antilles; New Caledonia;
New Zealand; Nicaragua; Nigeria; Niue; Northern Mariana Islands;
Oman; Pakistan; Palau; Panama; Papua New Guinea; Peru; Philippines;
Pitcairn; Portugal; Puerto Rico; Qatar; Saint Helena; Saint Kitts
and Nevis; Saint Lucia; Saint Pierre and Miquelon; Saint Vincent
and the Grenadines; Samoa; Sao Tomé and Principe; Senegal;
Sierra Leone; Singapore; Solomon Islands; Somalia; South Africa;
Spain; Sri Lanka; Suriname; Taiwan, Province of China; Tanzania,
United Republic of; Thailand; Timor-Leste; Togo; Tonga; Trinidad
and Tobago; Turks and Caicos Islands; United Arab Emirates; United
Kingdom; USA; Uruguay; Venezuela; Viet Nam; Virgin Islands, British;
Virgin Islands, U.S.; Wallis and Futuna; Western Sahara; Yemen.
S. coeruleoalba is categorised as "Least Concern"
by the IUCN (Hammond et al. 2008). The species is listed in Appendix
II of CITES. The eastern tropical Pacific population and the Mediterranean
populations are included in Appendix II of CMS. However, observations
off the coast off Japan also indicate migratory behaviour in these
waters. Range states concerned in these waters are Japan, North
and South Korea, the Peoples Republic of China and Taiwan (see Perrin
et al. 1996 in Appendix
2). Therefore, it is recommended that the West Pacific Stock
also be included in Appendix II of CMS.
To date, striped dolphins have faced relatively few threats compared
with other small cetacean species, although very little is known
about the species in some areas. However, some discrete populations
are affected either by both direct and indirect catches or by habitat
encroachment. In particular the direct catches off the Pacific coast
of Japan are a matter of concern, as was expressed by the International
Whaling Commission. The levels of contamination in the Mediterranean
Sea, coupled with the increasing incidental catches in the driftnet
fishery and reduced prey availability represent the major threats
for this and other cetacean species in the area.
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