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Globicephala melas (Traill, 1809) 
English: Long-finned pilot whale
German: Grindwal
Spanish: Calderón negro
French: Globicéphale noir
Drawing of Globicephala melas © Würtz-Artescienza (see
"links")
1. Description
The body in pilot whales is robust, with a thick 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 (Olson and Reily, 2002).
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 and Reily, 2002).
2. Distribution
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 Acores, Madeira, and Mauritania, including the
western Mediterranean (Rice, 1998 and refs. therein).
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).
Distribution of Globicephala melas (mod. from
Olson and Reilly 2002; © CMS / GROMS):
"antitropical" in cold temperate and subpolar waters of
all oceans except the North Pacific
(Carwardine, 1995). Enlarge
map.
Distribution of Globicephala melas in UK and
adjacent northern European waters
(Maps are interpreted from Sea Watch Foundation data and knowledge
of the ecology
of the species; from Evans 1998) Status: Regular, common or fairly
common (dark
shading); Regular, uncommon (fairly dark shading); Occasional (intermediate
shading);
Casual/Absent (light shading). Enlarge
map
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) summarise 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).
3. Population size
There is little information on stocks within the species. 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
(CV=0.295) by Buckland et al. (1993). 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). Canadas and
Sagarminaga (2000) report on observations in the the Alboran Sea,
an important oceanographic transition zone between the Mediterranean
and the Atlantic Ocean. Between April and September 1992-1997, the
authors sighted 109 pods. Comparison of results for encounter rate
and group size with those for other Mediterranean regions, together
with site fidelity shown by photo-identification and observations
of reproductive behaviour, emphasise the Alboran Sea as being one
of the most important areas for this species in the Mediterranean.
The average depth at encounters was 848.7 + 281.2 m ranging from
300 to 1,800 m, and reflecting the distribution of their preferred
diet, pelagic cephalopods.
Around the Faroe Islands pilot whales occur all year round with
a peak abundance in July-September. New tracking studies show a
preference over the border of the continental shelf (Bloch et al.,
1993c; Bloch et al., 2003).
Off the coast of Chile, Aguayo et al. (1998) mainly sighted G.
melas in proximity of the coast, reflecting its preference for
the edge of the continental shelf.
Goodall and Macnie (1998) report 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 south-western
South Atlantic, sightings clustered in two areas, 34- 46°S and
off Tierra del Fuego, 52-56°S. Here schools were found up to
1,000 nm 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.
Behaviour: Entire pods can sometimes be seen logging, allowing
close approach by boats. The strong blow may be visible in calm
weather (Carwardine, 1995).
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; Canadas 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) describe 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).
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 ans 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) report 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 down to 828 m depth
(Carwardine, 1995, Heide-Jørgensen et al., 2003). 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 Northeast United States, however, Atlantic mackerel (Scomber
scombrus) is said to be an important prey item, at least
during winter and early spring (Abend and Smith, 1997). Olson and
Reilly (2002) summarize that the diet in the northwest Atlantic
includes cod (Gadus morhua), turbot (Scomber
scombris), herring (Clupea harengus), hake
(Merluccios bilinearis; Urophysis spec.) and dogfish (Squalus
acanthias). The squid Todarodes sagittattus and species
of the genus Gonatus are reported prey items of long-finned
pilot whales in the eastern North Atlantic. 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). Werth (2000) describes the
feeding mechanism in captive juvenile long-finned pilot whales:
Depression and retraction of the large, piston-like tongue generate
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; similar observations on relative abundance of pilot
whales and squid are reported from the Faroe Islands (Reyes, 1991
and ref. therein; Bernard and Reilly, 1999 and refs. therein). According
to Carwardine (1995) both subspecies prefer deep water. While some
live permanently offshore or inshore, others make inshore (summer
and autumn) to offshore (winter and spring) migrations.
Mate (1989) tracked a pilot whale with an Argos satellite-monitored
radio tag for 95 days in the western North Atlantic. The whale was
located by satellite during movements of at least 7,588 km and sighted
from an aircraft several times in the company of other pilot whales.
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.
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. A catch statistics excist from the Faroes since 1584, unbroken
from 1709-today, showing an annual average catch of 850 pilot whales
(range: 0 - 4,480) with a cyclic variation according to the North-Atlantic
climatic variations (Bloch and Lastein, 1995; Bloch, 1998). In Greenland,
fisheries are minor (Butterworth, 1996).
Incidental catch: Incidental catches are reported from Newfoundland,
the Mediterranean and the Atlantic coast of France. In British waters,
long-finned pilot whales are accidentally caught in gillnets, purse
seines and in trawl fisheries. Very few are reported taken incidentally
in fisheries in the southern hemisphere (Reyes, 1991 and refs. therein).
However, 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 anomalously high take. 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).
Zerbini and Kotas (1998) report 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 Globicephala 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 squids are widespread
in the western North Atlantic. Target species for these fisheries
are squids eaten by pilot whales, making these vulnerable to prey
depletion.
Pollution: Long-finned pilot whales off the Faroes, France,
UK and the eastern USA appear to be carrying high levels of DDT
and PCB in their tissues. However, those animals examined off the
Newfoundland and Tasmanian coasts had very low levels, at least
of DDT. 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).
A different form of pollution has recently been investigated by
Rendell and Gordon (1999): The increasing level of man-made noise
in the world's oceans may have an effect on acoustically sensitive
groups such as cetaceans. 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.
7. Remarks
The only current fishery for long-finned pilot whales is undertaken
in the Faroe islands and Greenland. 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.
As well as the IWC, ICES and NAMMCO have concluded, that with an
estimated population size of 778 000 (CV=0.295) 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 also
reduces any threat.
Globicephala melas is not listed by the IUCN (see "links").
The North and Baltic Sea populations have been listed in Appendix
II of CMS. However, recent data on 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 that migrating between United States 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)
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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Kindly reviewed by Dorete Bloch, Museum of Natural History,
Thorshavn, Faroe Islands
©
Boris Culik, Kiel, Germany, 2003
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