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Globicephala melas (Traill,
1809) 
English: Long-finned pilot whale
German: Langflossen-Grindwal
Spanish: Calderón negro
French: Globicéphale noir
Family Delphinidae
Globicephala melas © Würtz-Artescienza (see "links")
1. Description
The body in pilot whales is robust, with a deep tail stock. The
melon is exaggerated and bulbous and the beak is barely discernible
or non-existent. The dorsal fin is wide, broad based, falcate and
set well forward on the body. The flippers are long, slender, and
sickle-shaped. A faint grey saddle patch may be visible behind the
dorsal fin in southern Hemisphere specimens. In the North Atlantic,
a thin whitish stripe can be visible in less than half of all adult
pilot whales. A pale eye blaze is visible in one fifth of all adult
pilot whales, most often in males (Bloch et al. 1993a). A grey midventral
line extends to the front into an anchor-shaped chest patch and
widens posteriorily to a genital patch. Sexual dimorphism exists
with longer flippers and larger flukes in males (Bloch et al. 1993a).
The long-finned pilot whale has a narrower skull than the short-finned
species, with the maxillary bones exposed laterally along the full
length of the rostrum (Olson, 2009).
Long- and short-finned pilot whales (G. melas and G. macrorhynchus)
are difficult to distinguish at sea. However, the species differ,
as the name suggests, in flipper length, skull shape and number
of teeth. On average, the flippers reach 18-30% of the body length
in long-finned pilot whales, but only 14-19% in short-finned pilot
whales (Bloch et al. 1993a). Adults reach a body length of approx.
6.5 m, males being 1 m larger than adult females (Bloch et al. 1993b;
Olson, 2009). Body mass reaches up to 1,300 kg in females and up
to 2,300 kg in males (Jefferson et al. 2008). 
2. Distribution
Two subspecies are recognized in some classifications (Rice, 1998):
G. m. melas: This subspecies ranges in the North Atlantic
from Ungava Bay, Disko in western Greenland, 68°N in eastern
Greenland, Iceland, the Faroes, and Nordland in Norway, south to
North Carolina, the Azores, Madeira, and Mauritania, including the
western Mediterranean (Rice, 1998 and refs. therein). It occurred
as recently as the 8th to 12th century in northern Japanese waters
(Olson, 2009).
According to Bloch and Lastein (1993) pilot whales on the western
(Newfoundland) and eastern (Faroes) sides of the North Atlantic
are distinguishable by minor external morphometric characters and
may be geographically isolated from each other. However, Fullard
et al. (2000) concluded that despite genetic, morphometric, physiological
and observational studies, it remains unclear whether any population
substructure exists. They used eight highly polymorphic microsatellite
loci to analyse samples from the US East Coast (Cape Cod), West
Greenland, the Faeroe Islands and the UK. Although their results
indicate that substructure does exist, and is particularly pronounced
between West Greenland and other sites, the magnitudes of the various
pairwise comparisons do not support a simple isolation-by-distance
model. Instead, the patterns of genetic differentiation suggest
that population isolation occurs between areas of the ocean which
differ in sea surface temperature (Fullard et al. 2000).
Distribution of Globicephala melas: the
species is "antitropical" in cold temperate and subpolar
waters of all oceans except the North Pacific (Taylor et al. 2008;
© IUCN; click
here for large map)
G. m. edwardii (A. Smith, 1834): This subspecies
is circumglobal in the Southern Hemisphere, ranging north to São
Paulo in Brazil, Cape Province in South Africa, Iles Crozet, Heard
Island, the southern coast of Australia, Great Barrier Island in
New Zealand, and Arica (19°S) in Chile. Southward it extends
at least as far as the Antarctic Convergence 47°S to 62°S
and has been recorded near Scott Island (67°S, 179°W) and
in the central Pacific sector at 68°S, 120°W (Rice, 1998
and refs. therein).
3. Population size
There is little information on stocks within the species, and there
is no information on global trends in abundance (Taylor et al.,
2008). The best northwestern Atlantic abundance estimate for Globicephala
sp. covering most of the species' habitat stems from two 2004 U.S.
Atlantic surveys: 31,139 whales (CV= 0,27; Waring et al,. 2007).
Based on surveys in the 1980's there are about 13,000 short-finned
pilot whales off eastern Newfoundland. In the north-eastern Atlantic
the number of pilot whales inhabiting the area between East Greenland,
Iceland, Jan Mayen, Faroe Islands and off the western coasts of
the British Islands and Ireland was estimated at around 778,000
by Buckland et al. (1993). However, in a more recent meeting, the
North Atlantic Marine Mammal commission (NAMMCO, 2006) noted that
there had been no assessment of pilot whales since 1994.
Estimates for Antarctic waters are in the order of 200,000 long-finned
pilot whales (Bernard and Reilly, 1999 and refs. therein).
4. Biology and Behaviour
Habitat: The typical temperature range for the species is
0-25°C (Martin, 1994) and it may be found in inshore but mostly
in offshore waters (Reyes, 1991 and refs. therein).
Around the Faroe Islands, pilot whales show a preference for the
region over the border of the continental shelf (Bloch et al. 1993c;
Bloch et al. 2003). In the the Alboran Sea, between the Mediterranean
and the Atlantic Ocean, the average depth of encounters was 849
m, ranging from 300 to 1,800m, and reflecting the distribution of
their preferred diet, pelagic cephalopods (Canadas and Sagarminaga,
2000).
In the southern hemisphere, off the coast of Chile, Aguayo et al.
(1998) mainly sighted G. melas close to the coast, reflecting its
preference for the edge of the continental shelf. Goodall and Macnie
(1998) reported on sightings in the south-eastern South Pacific,
which were clustered from 30-35°S, 72-78°W, the maximum
being about 160 nm from shore.
In the southwestern South Atlantic, sightings clustered in two areas,
34-46°S and off Tierra del Fuego, 52-56°S. Here schools
were found up to 1,000nm from shore. Fifteen sightings were from
waters south of the Antarctic Convergence, from December to March.
Only one sighting was made south of 44°S in winter, probably
due to lack of effort in southern seas during the colder months
(Goodall and Macnie 1998).
Behaviour: Mate (1989) tracked a pilot whale with an Argos
satellite-monitored radio tag for 95 days in the western North Atlantic.
Virtually all deep dives occurred at night, when the whale was likely
feeding on squid. Surface resting occurred most often immediately
after sunrise on a four- to seven-day cycle.
Bloch et al. (2003) tagged three long-finned pilot whales off the
Faroe Islands (62 °N, 7°W) with satellite transmitters.
After tagging, the whales separated and went in different directions.
After 10 days, two of the whales were observed together in a pod,
and after 19 days two were located at positions determined to be
within 2.3 km of each other. The swimming speed of the whales was
estimated at 0.2-14.5 km/hour, and they travelled average distances
of 70-111 km with a maximum of 200 km per day.
Baird et al. (2002) radio-tagged 5 long-finned pilot whales in deep
(>2000 m) waters of the Ligurian Sea, off the NW coast of Italy.
During the day all 5 whales spent their time in the top 16 m of
the water column, and visible surface activities consisted primarily
of rest and social behaviour. Shortly after sunset two whales made
several deep dives (max. 360 and 648 m) at high velocity, at a time
when vertically migrating prey become more readily available.
Schooling: Pilot whales are highly social; they are generally
found in pods of 110, but some groups contain up to 1,200 individuals
(Zachariassen, 1993; Bloch, 1998). Based on photo-identification
and genetic work, pilot whales appear to live in relatively stable
pods like those of killer whales, and not in fluid groups characteristic
of many smaller dolphins (Jefferson et al. 1993; Cañadas
and Sagarminaga, 2000). They are social animals, with close matrilineal
associations with 60% females.
The pods are often mixed with Atlantic white-sided dolphins (Lagenorhynchus
acutus) and bottlenose dolphins (Tursiops truncatus)
(Bloch et al. 1993c). When travelling, pods may swim abreast in
a line several kilometres across. Short-finned pilot whales are
often found in the company of bottlenose dolphins and other small
cetaceans, although they have been known to attack them (Carwardine,
1995). Baraff and Asmutis (1998) described the association of an
individually identified long-finned pilot whale with Atlantic white-sided
dolphins over six consecutive years. Pilot whales were also observed
in close association with fin, sperm and minke whales, and common,
bottlenose, hourglass and possibly dusky dolphins (Goodall and Macnie,
1998).
Off the northwest coast of Nova Scotia, Canada, Otttensmeyer and
Whitehead (2003) distinguished individuals on the basis of distinctive
marks on the dorsal fin. Animals formed short-term associations
over hours to days and long-term associations over years. Jankowski
(2006) found that groups consisted of 14.5 whales on average, with
a typical group size of 23 whales, which largely disassociates into
its component units (typically 8 whales) within a day. Within-unit
relationships typically lasted 4 years, but this is likely an underestimate.
G. melas is one of the species most often involved in mass
strandings, e.g. on Cape Cod (Massachusetts, USA) beaches from October
to January. Their tight social structure also makes pilot whales
vulnerable to herding, and this has been taken advantage of by whalers
in drive fisheries off Newfoundland, the Faroe Islands, and elsewhere
(Jefferson et al., 1993). If a whale of extreme social importance
or strong filial bond strands due to pathological or navigational
problems, others in the pod may strand also and then be unable to
remain off the beach once removed due to a secondary social or "caring"
response. This social response, however, was used successfully to
keep a pod of long-finned pilot whales from repeated strandings
by researchers in New Zealand: Because the "distress calls"
of the beached young of the pod appeared to evoke a stranding response
from the older whales, the younger whales were towed offshore and
moored to buoys, an action which lured the older animals back out
to sea (Bernard and Reilly, 1999 and refs. therein).
Reproduction: Mating occurs primarily in May-June and again
at a lower rate in October in the North Atlantic (Desportes et al.
1993; Martin and Rothery, 1993). Calving and breeding can apparently
occur at any time of the year, but peaks occur in summer in both
hemispheres (Jefferson et al. 1993).
Goodall and Macnie (1998) reported that young were present in all
areas of the south Pacific and South Atlantic, including the sub-Antarctic,
where they were seen in January (summer), March and April (autumn)
and October (spring), when a birth occurred, and in the Antarctic
in summer, with a birth occurring at South Georgia in March (autumn).
Food: Primarily squid eaters, pilot whales will also take
small medium-sized gregarious fish, when available (Desportes and
Mouritsen, 1993; Jefferson et al. 1993). They feed mostly at night,
when dives may last for 18 minutes or more and reach 828m depth
(Carwardine, 1995, Heide-Jørgensen et al. 2002). In the western
North Atlantic the main prey is the squid Illex illecebrosus, although
cod (Gadus morhua) or Greenland turbot (Rheinhardtius hippoglossoides)
may be eaten when squid is not available. Off the northeastern United
States, Atlantic mackerel (Scomber scombrus) is thought to
be an important prey item, at least during winter and early spring
(Abend and Smith, 1997). Olson (2009) described the diet in the
northwest Atlantic; it includes cod (Gadus morhua), turbot
(Scomber scombris), herring (Clupea harengus), hake
(Merluccios bilinearis; Urophysis spec.) and dogfish (Squalus
acanthias). Mintzer et al. (2008), however, found that long-finned
pilot whales off North Carolina feed predominantly on the long-finned
squid (Loligo pealei).
The squid Todarodes sagittatus and species of the genus Gonatus
are reported prey items of long-finned pilot whales in the eastern
North Atlantic (Olson, 2009). Although squids are the predominant
prey around the Faroe Islands, some fish, such as Argentina silus
and Micromesistius poutassou, are taken too. The whales in
this region do not appear to select cod, herring or mackerel, although
they are periodically abundant (Reyes, 1991 and refs. therein; Desportes
and Mouritsen, 1993; Bernhard and Reilly, 1999 and refs. therein).
Off the South Island of New Zealand, longfinned pilot whales feed
exclusively on cephalopods, mainly arrow squid, Nototodarus
spp., and common octopus, Pinnoctopus cordiformis (Beatson
and O'Shea, 2009).
Werth (2000) described the feeding mechanism in captive juvenile
long-finned pilot whales: Depression and retraction of the large,
piston-like tongue generated negative intraoral pressures for prey
capture and ingestion. Food was normally ingested without grasping
by teeth, yet was manipulated with lingual, hyoid, and mandibular
movement for realignment; suction was then used to transport prey
into the oropharynx.
5. Migration
In the Northwest Atlantic, pilot whales move towards the shelf
edge during mid winter through early spring, then move northward
along the edge to George's Bank and Nova Scotia, arriving off Newfoundland
in summer. The peak of the breeding season is said to be in August
in Newfoundland waters, where the whales remain until late autumn.
The inshore-offshore movements of pilot whales in the western North
Atlantic have been correlated with movements of their preferred
prey, squid (Reyes, 1991 and ref. therein; Bernard and Reilly, 1999
and refs. therein). Jankowski (2006) used photo-identification data
from two study sites 40 km apart, off the northwest coast of Nova
Scotia, Canada, between 1998 and 2003, to investigate habitat utilisation.
An average individual visits the study area for 1-2 days but may
return over a 5 year period. Individuals are well-mixed between
the two study sites. A satellite-monitored whale tracked for 95
days in the western North Atlantic was located during movements
of at least 7,588 km and sighted from an aircraft several times
in the company of other pilot whales (Mate 1989).
Site fidelity is also reported from the the Faroe Islands, where
pilot whales occur all year round with a peak abundance in July-September.
New tracking studies show a preference for the area over the border
of the continental shelf (Bloch et al. 1993c; Bloch et al. 2003).
6. Threats
Direct catch: Drive fisheries for long-finned pilot whales
in the Faroe Islands date back to the Norse settlement in the 9th
century. Catch statistics exist from the Faroes since 1584, unbroken
from 1709-today and showing an annual average catch of 850 pilot
whales (range: 0 - 4,480) with a cyclic variation correlated with
North-Atlantic climatic variations (Bloch and Lastein, 1995; Bloch,
1998). Considering the mobility of these animals, it seems likely
that these catches are recruited from a larger area in the North
Atlantic than previously assumed. This suggests that the whales
are taken from a larger population than that estimated from coastal
areas around the Faroe Islands, hence increasing the probability
that the harvest is sustainable (Bloch et al. 2003). In Greenland,
fisheries are minor (Butterworth, 1996).
Incidental catch: Incidental catches are reported from Newfoundland,
the Mediterranean and the Atlantic coast of France, and according
to Bernard and Reily (1999 and refs. therein) there are probably
more pilot whales taken incidentally than are presently documented.
On the east coast of the USA, the foreign Atlantic mackerel fishery
was responsible for the take of 141 pilot whales in 1988. This fishery
was suspended in early May of that year as a direct result of this
high take. More recently, most of the estimated marine mammal bycatch
is from U.S. Atlantic EEZ waters between South Carolina and Cape
Cod in the pelagic longline fishery. The average annual Globicephala
spp. mortality in 2000-2004 was 70 animals (CV=0.37). The average
annual estimated fishery-related mortality in the northeast mid-water
trawl fishery during 2002-2004 was 8.9 (CV= 0.35) (Waring et al.
2007).
In British waters, long-finned pilot whales are accidentally caught
in gillnets, purse seines and in trawl fisheries (Reyes, 1991 and
refs. therein). The seas around Cornwall, SW Britain, are one of
the most heavily fished areas of the UK, and Leeney et al. (2008)
found that strandings of pilot whales around Cornwall have increased
significantly since the mid-1970s, with seasonal peaks in stranding
frequencies between November and January. Sixty-one % of investigated
individuals were determined to have died due to bycatch in fishing
gear.
A 1990 workshop to review mortality of cetaceans in passive nets
and traps documented an annual kill of 50-100 G. melas off the Atlantic
coast of France. Furthermore, pilot whales are also known to be
taken incidentally in trawl and gillnet fisheries in the western
North Atlantic and in swordfish driftnets in the Mediterranean (Jefferson
et al. 1993). This seems to be still ongoing, as Lopez et al. (2003)
report that around 200 cetaceans might be caught annually in inshore
waters and around 1500 in offshore waters of Galicia (NW Spain),
mainly small dolphins, as well as Tursiops truncatus and Globicephala
melas.
Very few were reported taken incidentally in fisheries in the southern
hemisphere (Reyes, 1991 and refs. therein). However, Zerbini and
Kotas (1998) reported on cetacean-fishery interactions off southern
Brazil. The pelagic driftnet fishery is focused on sharks (families
Sphyrnidae and Carcharinidae) and incidentally caught species include
15 G. melas in 1995 and 1997. Authors conclude that the driftnet
fishery may be an important cause of cetacean mortality and that
a systematic study should be carried out in order to evaluate the
impact of this activity.
Overfishing: Commercial fisheries for squid are widespread
in the western North Atlantic. Target species for these fisheries
are squid species which form a large part of the diet of pilot whales,
making these vulnerable to prey depletion (Taylor et al. 2008).
Ship strikes: Since high speed ferries were introduced in
the Canary Islands in 1999, their number has grown steadily, and
collisions with cetaceans have been reported ever since. While true
numbers of collisions remain unknown, estimates range from approx.
10 to 30 cetaceans killed every year. Present knowledge indicates
that the sperm whale is the species most frequently hit, but baleen,
beaked and pilot whales are affected as well (Weinrich et al. 2005)
Pollution: Long-finned pilot whales off the Faroes, France,
UK and the eastern US appear to be carrying high levels of DDT and
PCB in their tissues, and where whales are consumed by humans, this
leads to high-level burdens of organohalogens among residents, e.g.
at the Faroes (Faengstroem et al. 2005). In other parts of their
distributional range, e.g. off Newfoundland and Tasmania, very low
levels of DDT were detected. Heavy metals such as cadmium and mercury
also have been found in pilot whales from the Faroes. Because these
contaminants accumulate in tissues over time, older animals and
especially adult males tend to have higher concentrations (Borell
and Aguilar, 1993; Caurant et al. 1993; Caurant and Amiard-Triquet,
1995). Combinations and levels of these pollutants may one day play
a role in stock differentiation (Reyes, 1991 and refs. therein;
Bernard and Reilly, 1999 and refs. therein; Frodello et al. 2000;
Nielsen et al. 2000).
Weisbrod et al. (2000) characterised organochlorine bioaccumulation
in pilot whales collected from strandings in Massachusetts and caught
in nets. Whales that stranded together had more similar tissue-levels
than animals of the same gender or maturity, reflecting pod-fidelity.
The high variation in tissue concentrations among individuals and
pods, and the similarity within a stranding group suggest that pilot
whale pods are exposed to a large range of pollutant sources, such
as through different prey and feeding locations (Desportes et al.
1994).
Noise pollution: The military makes extensive use of underwater
sound in order to find targets such as ships and submarines, and
some active military sonar systems are known to use very loud sources.
However, in part because these systems are classified, the characteristics
of such sound sources have rarely been published, and there have
been few studies of their effects on cetaceans. Although Rendell
and Gordon (1999) could not show any deleterious consequences for
the species, recordings of vocalisations indicated short-term vocal
responses of long-finned pilot whales to the sound source. However,
in 2005, three mass stranding events occurred in Tasmania, Australia,
involving approximately 145 long-finned pilot whales. The first
occurred six hours before the arrival of two Royal Australia naval
vessels, the second event began just over an hour after the vessels
began using high frequency (50-200 kHz) sonar in the vicinity of
the stranding. A behavioral reaction to the sonar facilitating the
second and third stranding events could not be ruled out (Parsons
et al. 2008).
7. Remarks
Range states (Taylor et al. 2008):
Algeria; Argentina; Australia; Belgium; Brazil; Canada; Chile; Denmark;
Falkland Islands (Malvinas); Faroe Islands; France; French Southern
Territories (the) (Crozet Is.); Germany; Greenland; Heard Island
and McDonald Islands; Iceland; Ireland; Italy; Libyan Arab Jamahiriya;
Malta; Morocco; Namibia; Netherlands; New Zealand (Antipodean Is.,
Chatham Is., North Is., South Is.); Norway; Peru; Portugal (Azores,
Madeira); South Africa (Marion-Prince Edward Is., Western Cape Province);
South Georgia and the South Sandwich Islands; Spain; Sweden; Tunisia;
United Kingdom; United States (North Carolina); Uruguay.
The only current fishery for long-finned pilot whales is undertaken
in the Faroe Islands. Although this fishery has been actively pursued
since the 9th century, catch levels have not shown evidence of depletion
of the stock as occurred off Newfoundland. ICES and NAMMCO as well
as the IWC, have concluded that with an estimated population size
of 778,000 in the eastern North Atlantic and approximately 100,000
around the Faroes (Buckland et al. 1993; NAMMCO, 1997) the Faroese
catch will not deplete the population. Pilot whales seem to utilise
a larger area around the Faroes (Desportes et al. 1994; Bloch et
al. 2003), which according to these sources also reduces any threat.
Globicephala melas is considered as "Data Deficient"
by the IUCN. The species is listed on CITES Appendix II.
The North and Baltic Sea populations have been listed in Appendix
II of CMS. However, data on long-range movements in the NW and NE
Atlantic suggest that these stocks should also be included in App.
II of CMS. Range states concerned are the US, Canada, Greenland,
Iceland, Norway, Ireland and the UK.
Attention should also be paid to the western North Atlantic population(s),
in particular migration between US and Canadian waters, formerly
depleted by overhunting and now facing increasing incidental mortality
in trawl fisheries (Reyes, 1991 and refs. therein).
As noted above, pollution (including noise pollution) by-catch and
mass strandings may be a threat to the species and warrant further
investigation. Population size and migratory patterns, including
home-range sizes are insufficiently known. For recommendations on
South American stocks, please see Hucke-Gaete (2000) in Appendix
1.
Please also see a report on the long-finned pilot whale posted on
the web by the North Atlantic Marine Mammal Commission: http://www.nammco.no.
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© Maps by IUCN.
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