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Physeter macrocephalus Linnaeus,
1758
English: Sperm whale
German: Pottwal
Spanish: Cachalote
French: Cachalot
Family: Physeteridae
Physeter macrocephalus © Wurtz-Artescienza
(see links)
1. Description
The sperm whale is the largest of the toothed whales, and there
is a strong sexual dimorphism between males and females: adult females
reach approx. 11 m in length and a body mass of 15,000 kg, whereas
males are much larger with 16 - 18 m body length and 45,000 - 57,000
kg (Whitehead, 2009, Jefferson et al. 2008). The most striking feature
of this species is its massive head, which makes up 25 - 30 % of
total length. It contains the spermaceti organ and the underlying
"junk", both set above the upper jaw and in front of the
parabolic-shaped facial region of the skull. These structures are
composed of spongy, oil-filled tissue enclosed in a muscular case
and bounded at both ends by air sacs. This spermaceti organ is responsible
for the striking echo-locating capabilities of sperm whales (Whitehead,
2009).
The single blowhole is set at the front of the head, offset to
the left and S-shaped. There is a thick, low dorsal hump instead
of a fin, about 2/3 back from the tip of the head, followed by a
series of bumps on the dorsal ridge towards the fluke. The flukes
are broad and triangular, resembling a ginkgo-leaf, with an almost
straight trailing edge showing individually different nicks, notches
and cuts, which are used in photo-identification. The body surface
is smooth over the head and wrinkled behind and on the sides. The
flippers are short, spatulate and wide. The lower jaw is narrow
and underslung and carries 18 - 26 pairs of functional teeth, which
are absent from the upper jaw. The predominant colour is black to
dark brownish-grey with white areas around the mouth and often on
the belly. Some animals have white to yellowish calluses on the
dorsal hump, and some adult males have white scratches on the head.
Albino sperm whales have been observed and one is well-known from
the world literature (Moby Dick, by Herman Melville). The characteristic
bushy blow is up to 5 m high and projects to the left of the animal
at an angle from the vertical (Jefferson et al. 2008).
2. Distribution
The sperm whale is one of the animals with the widest distribution
on the globe. It ranges from the ice-edge of both hemispheres as
far south as the equator but concentrates in so-called "grounds"
which coincide with areas of high marine productivity. Such areas
can measure a few hundred kilometres across and may contain hundreds
or even thousands of sperm whales (Rice 1989; Whitehead, 2009).
.
The sperm whale has a world wide distribution,
with females inhabiting the lower latitudes
below 40° (except in the north Pacific where they range to 50°N)
and males ranging north
and south of this boundary to the ice-edge of the Arctic and Antarctic
Oceans
(Whitehead, 2009; Taylor et al. 2008; © IUCN; enlarge
map)
Both sexes have very distinct distributions when mature.
Most females range usually far from land, except for areas near
seamounts or oceanic islands, in waters 1000 m deep or deeper. They
prefer water temperatures above 15 °C and low latitudes within
40°N and 40°S (except for the North Pacific where they range
to 50°N). Young males accompany these groups of females until
they are 4 - 21 years old and then migrate to higher latitudes.
There they may range to the ice-edge, preferring productive waters,
e.g. in the vicinity of deep sea canyons such as off Andenes, northern
Norway, or Kaikoura, New Zealand, but they may also be found in
more shallow waters e.g. off Nova Scotia or New York in water only
300 m deep (Whitehead, 2009).
Recent genetic studies have shown significant subdivision between
the Gulf of Mexico, the Mediterranean, and the North Atlantic (Drouot,
2003; Engelhaupt, 2004). Mitochondrial DNA results showed significant
differentiation among all populations, while microsatellites showed
significant differentiation only for comparisons with the Mediterranean
Sea, and at a much lower level than seen for mtDNA (Engelhaupt et
al. 2009).
3. Population size
Whitehead (2002) extrapolated from about one quarter of the global
habitat that the current population size world-wide is about 360,000
animals (CV = 0.36) or about 29% of the pre-whaling population size.
With a maximum rate of increase of around 1% per year (Whitehead
2002), the species is not well adapted to recover from population
depletion and the threats exposed below are made responsible for
the fact that regional populations of sperm whales are declining
or are apparently not recovering from depletion.
There have been several population surveys within the past 10-12
years:.
In the Northwest Pacific a preliminary analysis (Kato and Miyashita,
1998) indicated 102,112 (CV = 0.155) sperm whales, but this was
reduced to 29,674 (CV = 0.14) in Kato and Miyashta (2000) using
only primary sightings. In the eastern temperate North Pacific a
spring 1997 line-transect survey resulted in estimates of 24,000
(CV=0.46) individuals based on visual sightings and 39,200 (CV=0.60)
based on acoustic detections and visual group size estimates (Barlow
and Taylor 1998). In the EEZ waters off the US west coast from Washington
to Califiornia, the most recent of abundance estimate is 2,853 (CV=0.25)
animals from ship surveys conducted in 2001 (Barlow and Forney 2007)
and 2005 (Forney 2007). In the eastern tropical Pacific, abundance
was estimated as 22,700 (95% C.I.=14,800-34,600; Wade and Gerrodette
1993) and in US EEZ around the Hawaii islands, a 2002 shipboard
line-transect survey resulted in an abundance estimate of 7,082
(CV=0.30) sperm whales (Barlow 2003), including a correction factor
for missed diving animals. No data on current population trends
are available from any of these regions (Angliss and Allen, 2008).
In the Northwest Atlantic two 2004 surveys yielded 4,804 (CV =0.38),
with an estimate from the northern U.S. Atlantic of 2,607 (CV =0.57)
and from the southern U.S. Atlantic of 2,197 (CV =0.47). In the
northern Gulf of Mexico oceanic waters, the abundance estimate pooled
from 2003 to 2004 is 1,665 (CV=0.20) (Mullin 2007). Similar to Pacific
estimates, there are insufficient data to determine population trends
(Waring et al. 2009). In the Northeast Atlantic the most recent
estimates are dated, with 6,013 (CV = 0.32) (Christensen et al.
1992) and 1,772 (CV= 0.18) in the Faroes-Iceland area (Gunnlaugsson
and Sigurjónsson, 1990).
Previous and historical population estimates from the Indian Ocean
and the southern Hemisphere are deemed unreliable and therefore
not presented here (NMFS, 2009).
4. Biology and Behaviour
Habitat: Sperm whales show a strong preference for deep
waters (Rice 1989), especially in areas with high sea floor relief.
Along the U.S. east coast, the overall distribution is centred along
the shelf break and over the continental slope 90-1,800 m deep (CETAP
1982; Waring et al. 2005). Very high densities occur in inner slope
waters north of Cape Hatteras, North Carolina seaward of the 1,000
m isobath during summer months (Waring et al. 2005). Sperm whales
are also known to move onto the continental shelf in waters less
than 100 m deep on the southern Scotian Shelf and south of New England,
particularly between late spring and autumn (Whitehead et al. 1992a
and b).
P. macrocephalus off Andenes, Norway © Boris
Culik
In the northern Gulf of Mexico, they occur in greatest density
along and seaward of the 1,000 m isobath (Mullin et al. 1991, Mullin
and Fulling 2004). They appear to prefer steep rather than shallow
depth gradients (Davis et al. 1998). The spatial distribution of
sperm whales within the Gulf is strongly correlated with mesoscale
physical features such as loop current eddies that locally increase
primary production and prey availability (Biggs et al. 2005). However,
sperm whales may also be found in rather unproductive marine areas
such as the "Charleston grounds" in the Sargasso Sea (Jacquet
1996). These are though to be associated with spawning squid (Jacquet
and Whitehead 1996).
Behaviour: On average, sperm whales spend more than 72%
of their time in foraging dive cycles. While foraging they make
repeated long dives, with a modal depth of 985 m in the Atlantic
Ocean, 644 m the Gulf of Mexico and 827 m the Ligurian Sea. Dive
durations of about 45 min are interrupted by surface intervals lasting
about 9 min (Watwood et al. 2006). However, dives can be much deeper
or shallower, and dive durations are also quite flexible (Whitehead,
2009). Dives are initiated by a deep breath, followed by a raising
of the head, submersion of the animal, and curving of the tail stock
above the water surface until the flukes are raised nearly vertically
out of the water. Descent to depth as well as return to the surface
can be nearly vertical. During the initial phase of the descent
the whales remain quiet, presumably navigating visually, but after
reaching a depth of between 100 - 220 m they initiate a series of
clicks emitted in intervals of 0,5 - 1 s. Watwood et al. (2006)
found that sperm whales descend a mean of 392 m from the start of
this regular clicking to the first buzz, i.e. accelerated clicks
associated with prey detection, which supports the hypothesis that
regular clicks function as a long-range biosonar.
During the late afternoon, females and young gather near the surface
to rest, in close aggregation. However, at the onset or end of foraging
activities, their behaviour may also show breaches, lobtails, manoeuvres,
rolls, and touching of conspecifics, while emitting codas, i.e.
stereotyped sequences of clicks, or creaks (Whitehead, 2009).
Schooling: Female sperm whales are organized in groups in
which adults travel with their sub-adult offspring. Males eventually
leave these groups, at age 4 to 21, after which they live in "bachelor
schools". The cohesion among males within a bachelor school
declines as the animals age (Best 1979) and they generally move
to higher latitudes. During their prime breeding period and old
age, male sperm whales are essentially solitary (Christal and Whitehead
1997).
Entire schools of sperm whales occasionally strand, but the causes
of this phenomenon are uncertain (Rice 1989). Changes in wind patterns
which result in colder and presumably nutrient-rich waters being
driven closer to the surface (Evans et al. 2005), lunar cycles (possibly
as a result of the effects that light levels have on the vertical
migration of their prey species (Wright 2005)), and solar driven
geomagnetic variations (Vanselow and Ricklefs 2005), which over
the last 400 years may account for 20% of all observed stranding
events around the North Sea (Vanselow et al. 2009), all play a role.
Reproduction: Females become sexually mature at 7-13 years
of age. The peak breeding season is generally in the spring: in
the northern hemisphere between March/April and June, and in the
southern hemisphere between October and December (Best et al. 1984).
Gestation lasts between14 - 16 months and females lactate for at
least two years. The inter-birth interval is 4-6 years for prime-aged
females. Female sperm whales rarely become pregnant after the age
of 40 (Best et al. 1984; Whitehead 2003).
Puberty in males usually begins between the ages of 10 and 20, and
most individuals do not become fully mature until their late twenties
(Best, 1979). Longevity can reach at least 50 years (Whitehead,
2009).
Food: Sperm whales forage in mesopelagic and benthic habitats,
primarily targeting cephalopods (Kawakami 1980) but occasionally
also fish (Clarke et al. 1993). Histioteuthids, mesopelagic gelatinous
squid ranging in mass between 0.1 and 1 kg, are at the top of the
list of preferred food items. However, females also feed on larger
prey such as giant squid (Archaeteuthys sp.) and jumbo squid
(Dosidicus sp.), and males prey on species such as the Antarctic
colossal squid (Mesonychoteuthis hamiltoni) (Whitehead, 2009).
The yearly turnover of biomass by sperm whales is estimated to be
comparable to the total catches of human fisheries, and the impact
of sperm whales on deep ocean food webs and nutrient cycling in
the ocean is probably significant (Clarke 1976; Kanwisher &
Ridgway 1983; Whitehead 2003).
Stomachs of male sperm whales from the Northeast Atlantic stranded
in 1997 on the Netherlands coast, in 1998 at Bettyhill (Scotland)
and in 1996 at Tory Island (Ireland) consisted almost entirely of
cephalopod beaks, with some containing also fish remains. Gonatus
sp. (probably Gonatus fabricii, Oegopsida: Gonatidae) was
the main prey. The fish remains were saithe (Pollachius virens,
Gadiformes: Gadidae), and remains of monkfish (Lophius sp.,
Lophiiformes: Lophiidae). A specimen stranded in Ireland had consumed
a wider range of prey, mainly Histioteuthis bonnellii (Oegopsida:
Histiotetuhidae), but also Architeuthis sp. (Oegopsida: Architeuthidae),
Chiroteuthis sp. (Oegopsida: Chiroteuthidae), Teuthowenia
megalops (Oegopsida: Cranchiidae) and the octopod Haliphron
atlanticus (Incirrata: Alloposidae) (Santos et al. 2002).
Cephalopod beaks from the stomachs of 10 males stranded in Denmark
between 1991 and 2000 revealed Gonatus fabricii as the dominant
prey species (>98%), the majority (73.5%) individuals with an
estimated mantle length between 192 and 257 mm. Other species found
were: Todarodes sagittatus, Histioteuthis sp., Cyclioteuthis
sp., Haliphron atlanticus, and Bathypolypus sp. (Simon
et al. 2003).
5. Migration
Movements and migratory behaviour of sperm whales has been studied
using photo-identification, tags, and satellite tags as well as
by following individual groups at sea. When feeding conditions are
adequate, sperm whales usually stay in small areas, 10 - 20 km across.
During travelling episodes the animals cover about 4 km/hr or about
90 km/day. Female home ranges are approx. 2,000 km across, but males
roam more widely (Whitehead, 2009).
Whitehead (2003) found that females and juveniles identified off
the Galápagos Islands frequently moved to the waters off
the mainland of Ecuador, more than 1,000 km away, to Panama and
Perú, about 1,500 - 2,000 km away and rarely to Chile or
California, travelling more than 3,000 km. Long-range movements
are also reported by tagged females in the South Pacific, where
females tagged for periods of over one month travelled on average
650 km (Best, 1979).
Available data suggests periodic migrations of mature males between
low latitude breeding and high latitude feeding grounds (Best, 1979).
Tagged males in the Southern Ocean showed average displacements
of 1,600 km (Best, 1979) or about twice the value recorded for females.
In the North Pacific, tagged whales were identified as having traveled1,300
km (Kasuya and Miyashita, 1988). There are more extreme values,
such as a male marked off Nova Scotia and killed off Spain, 4,300
km away (Mitchell, 1975), or a male marked south of Mexico and killed
off British Columbia, 4,850 km to the North (Kasuya and Miyashita,
1988) and finally a male marked off the north African coast and
killed off South Africa, 7,400 km to the South (Ivashin and Rovnin,
1967). In their feeding grounds, males off Kaikoura, New Zealand,
or Bleik Canyon, Andenes, Norway, show long term site fidelity,
returning over years to the same sites (Whitehead, 2003).
These findings are confirmed by more recent investigations. Genetic
comparison of putative sperm whale populations located in the Gulf
of Mexico, western North Atlantic, Mediterranean Sea and North Sea
indicate a strong fidelity of females to coastal basins on either
side of the North Atlantic and suggest the movement of males among
these populations for breeding (Engelhaupt et al. 2009). This confirms
previous genetic studies based on maternally inherited markers showing
inter-oceanic movement to be more prevalent among males than females
(Lyrholm et al. 1999) and is consistent with observation of females
having smaller geographic ranges.
6. Threats
Direct catch: Local whaling dates back to the 1500s and
intense commercial whaling to around 1712. Highly mechanised "modern"
whaling was particularly intense around 1950, and at its peak killed
around 25,000 whales per year, significantly depleting the global
population. After the cessation of commercial whaling in 1989, the
annual catch decreased to some tens of whales taken each year from
small boats in Indonesia (Reeves 2002), although none have been
taken in recent years (H. Whitehead, pers. comm. to Taylor et al.
2008), and 10 taken annually by Japan under a special IWC scientific
permit (Clapham et al. 2003).
Incidental catch: Incidental capture in fishing gear, such
as gillnets and bottom-set longline gear, continues to take a toll
on sperm whale populations, although the degree of threat is considered
low. They have been found as bycatch in pelagic drift gillnets targeting
swordfish and tuna in U.S. east-coast waters (Waring et al. 1997),
and in artisanal gillnets targeting sharks and large pelagic fishes
off the Pacific coasts of northwestern South America, Central America,
and Mexico (Palacios and Gerrodette 1996). The pelagic drift gillnet
fishery closed in 1997 and the use of drift gillnets was prohibited
in 1999, but sperm whales are still threatened by fishing gear.
An estimated average of >0.2 sperm whales are killed or seriously
injured annually in the driftnet fishery for thresher sharks and
swordfish and unknown fisheries off Oregon and California (Carretta
et al. 2009). No estimates of mortality are available for the Mexican
driftnet fisheries (NMFS, 2009).
In 2006, there were three observed serious injuries in the Gulf
of Alaska sablefish longline fishery, which extrapolates to 10 estimated
serious injuries for that fishery for that year (R.P. Angliss, in
NMFS, 2009). Entanglements in longline fishing gear have also been
observed in South Georgia (Purves et al. 2004) and Chile (Ashford
et al. 1996). Sperm whales have been found following deep-water
trawlers during hauling operations targeting Greenland halibut,
and one case of entanglement in the trawl was reported (Karpouzli
and Leaper 2004). Between 1998-2006, no sperm whales were known
to be killed due to fishery interactions in the U.S. Atlantic Gulf
of Mexico (Waring et al. 2009), indicating that current fishing
practices pose a low threat to the recovery of sperm whale populations.
However, unreported by-catch and levels of mortality and injury
due to entanglement in lost or discarded gear are still a matter
of concern, especially in the Mediterranean (Taylor et al. 2008).
Even in the absence of whaling, the Mediterranean population appears
to have declined over the past 20 years, with bycatch in driftnets
targeted at swordfish a likely principal cause (Reeves and Notarbartolo
di Sciara 2006).
Killing: Although the magnitude of these interactions is
infrequently documented, there are reports of sperm whales having
been killed to keep them away from fishing gear (Gonzalez 2001).
Pollution: Increases in the rate of sperm whale strandings
in western Europe since the early 1980s have raised concern that
anthropogenic effects, including pollution, may be a contributing
factor (Goold et al. 2002). However, the results of a tissue analyses
of stranded whales for a wide range of contaminants showed no clear
link between contamination and stranding (Jacques and Lambertsen
1997). Nevertheless, levels of mercury, cadmium, and certain organochlorines
in these whales' tissues were high enough to cause concern about
toxicity and other possibly indirect health effects (Bouquegneau
et al. 1997; Law et al. 1997). Fossi et al. (2003) stated that high
concentrations in the Mediterranean could have an effect on reproductive
rates of this species. The levels of organochlorine compounds found
in females were consistently higher than those in males, which is
contrary to the typical findings in other marine mammals. Given
that male and female sperm whales are geographically separated during
much of the year, it is possible that males feed in less polluted
waters or perhaps on less contaminated prey than females (NMFS,
2009).
Noise pollution: Sperm whales may be adversely affected
by anthropogenic noise causing permanent or temporary damage to
their hearing, masking biologically relevant sounds, or enticing
negative changes in behaviour. However, it is difficult to ascertain
the level of threat from anthropogenic sound sources with currently
available information (NMFS, 2009). During seismic experiments in
the northern Gulf of Mexico a whale carrying an acoustic tag moved
away from an operating seismic vessel once the seismic pulses were
received at roughly 137 dB re 1 µPa (Johnson et al. 2003).
As opposed to this, Davis et al. (2000) noted that sighting frequency
in the northern Gulf of Mexico did not differ significantly among
different acoustic levels. Off Nova Scotia an active seismic program
did not elicit any obvious changes in sperm whale distribution or
behaviour (McCall 1999). Offshore northern Norway, animals continued
to call when exposed to pulses from a distant seismic vessel of
up to 146 dB re 1 µPa peak-to-peak (Madsen et al. 2002). And
seismic work off Angola (Weir 2008) resulted in no difference in
encounter rates of sperm whales or obvious behavioural changes due
to air gun activity.
However, in United Kingdom waters sperm whales exhibited some changes
in behaviour in the presence of operating seismic vessels (Stone
2003). And in the Gulf of Mexico Jochens et al. (2008) found decreases
in foraging activity.
There have been no sperm whale strandings attributed to naval sonar.
However, there is some evidence of disruptions of clicking and behaviour
from sonars, pingers, and the Acoustic Thermometry of Ocean Climate
studies (summarised in NMFS, 2009).
Ship strikes: The most recent statistics, in the world-wide
large whale ship strike database (Jensen and Silber 2004), indicate
that from 1975 to 2002 out of a total record of 292 strikes of all
large whales, sperm whales were struck 17 times, 13 of which resulted
in death of the whale. Vessel types include Navy vessels, container/cargo
ships, whale-watching vessels, cruise ships, ferries, Coast Guard
vessels, and tankers. The estimates of serious injury or mortality
should be considered minimum values because many ship strikes go
either undetected or unreported for various reasons (NMFS, 2009).
The most severe injuries are caused by larger vessels (80 m or longer)
and vessels travelling at a speed of 14 knots or faster (Laist et
al. 2001).
Sperm whales spend long periods (typically up to 10 minutes; Jacquet
et al. 1998) "rafting" at the surface between deep dives.
Reports of ships colliding with sperm whales are frequent in the
Canary Islands, where ship traffic is heavy and the local density
of sperm whales relatively high (André et al. 1997). In the
North Atlantic, a merchant ship reported a strike in Block Canyon
in May 2000 (Waring et al. 2007), and from 2001-2003, one stranded
sperm whale was reported struck by a naval vessel and another by
a merchant vessel near Rhode Island (Waring et al. 2005). More recently
in the Pacific, two sperm whales were struck by a ship in 2005,
but it is not known if these ship strikes resulted in death or injury
(NMFS, 2009). In general, however, it does not appear that ship
strikes are a significant threat to sperm whale populations (Whitehead
2003).
Whale watching: In Kaikoura, New Zealand resident whales
respond to whale-watching activities with small changes in ventilation
and vocalization patterns. These changes may not be of biological
importance (Richter et al. 2006). However, transient whales, which
receive less whale-watching effort, respond differently, and usually
more strongly to whale-watching boats. They spend less time at the
surface and adjust their breathing intervals and acoustic behaviour
(Gordon et al. 1992).
Marine debris: Instances of stomach obstruction caused by
marine debris have been documented in sperm whales, but severity
of threat is considered low due to the small number of known cases.
In 1989, a necropsy on a stranded sperm whale indicated that its
death was caused by a stomach obstruction following accidental ingestion
of plastic bags and sheets in the Lavezzi Islands of the Tyrrhenian
Sea (Viale et al. 1992). In Iceland a necropsied animal had a lethal
obstruction of the gut with plastic marine debris (Lambertsen 1990).
The stomach contents of two sperm whales that stranded separately
in California (California Marine Mammal Stranding Database 2008,
in NMFS, 2009) included extensive amounts of netting from discarded
fishing nets; however, the cause of death was not determined.
7. Remarks
Range states (Taylor et al. 2008) :
Albania; Algeria; Angola; Antarctica; Antigua and Barbuda; Argentina;
Australia; Bahamas; Bangladesh; Barbados; Belgium; Belize; Benin;
Brazil; Brunei Darussalam; Cameroon; Canada; Cape Verde; Chile;
China; Colombia; Comoros; Costa Rica; Croatia; Cyprus; Denmark;
Djibouti; Dominica; Dominican Republic; Ecuador; Egypt; El Salvador;
Equatorial Guinea; Falkland Islands (Malvinas); Faroe Islands; Fiji;
France; Gabon; Gambia; Ghana; Gibraltar; Greece; Greenland; Grenada;
Guatemala; Guinea; Guinea-Bissau; Guyana; Haiti; Honduras; Iceland;
India; Indonesia; Iran, Islamic Republic of; Ireland; Israel; Italy;
Jamaica; Japan; Kenya; Kiribati; Korea, Democratic People's Republic
of; Korea, Republic of; Lebanon; Liberia; Libyan Arab Jamahiriya;
Madagascar; Malaysia; Maldives; Malta; Marshall Islands; Mauritania;
Mauritius; Mexico; Micronesia, Federated States of; Monaco; Morocco;
Mozambique; Namibia; Nauru; Netherlands; Netherlands Antilles; New
Zealand; Nicaragua; Nigeria; Niue; Norway; Oman; Pakistan; Palau;
Panama; Papua New Guinea; Peru; Philippines; Portugal; Russian Federation;
Saint Helena; Saint Kitts and Nevis; Saint Lucia; Saint Vincent
and the Grenadines; Samoa; Sao Tomé and Principe; Senegal;
Seychelles; Sierra Leone; Singapore; Slovenia; Solomon Islands;
Somalia; South Africa; Spain; Sri Lanka; Suriname; Syrian Arab Republic;
Taiwan, Province of China; Tanzania, United Republic of; Thailand;
Timor-Leste; Togo; Tonga; Trinidad and Tobago; Tunisia; Turkey;
Tuvalu; United Kingdom; USA; Uruguay; Vanuatu; Venezuela; Viet Nam;
Yemen.
Sperm whales are classified as Vulnerable by the IUCN. Whitehead
(2002) provided a model-based estimate of global trend giving a
6% probability for Endangered, a 54% probability of meeting the
Vulnerable category, and a 40% probability of falling into the Near
Threatened category.
P. macrocephalus is on Appendix I of CITES and Appendices I
and II of CMS.
8. Sources
· André M, Terada M, Watanabe Y (1997)
Sperm whale (Physeter macrocephalus) behavioural response
after the playback of artificial sounds. Rep Int Whal Commn 47:499-504.
· Angliss RP, Allen BM (2008) Sperm whale (Physeter macrocephalus):
North Pacific Stock NOAA-TM-AFSC-193
· Ashford JR, Rubilar PS, Martin AR (1996) Interactions between
cetaceans and longline fishery operations around South Georgia.
Mar Mamm Sci 12:452-457.
· Barlow J (2003) Cetacean abundance in Hawaiian waters during
summer/fall 2002. Admin Rep LJ-03-13. Southwest Fisheries Science
Center, National Marine Fisheries Service, La Jolla, CA
· Barlow J, Forney KA (2007) Abundance and population density
of cetaceans in the California Current ecosystem. Fishery Bulletin
105:509-526.
· Barlow J, Taylor BL (1998) Preliminary abundance of sperm
whales in the northeastern temperate Pacific estimated from a combined
visual and acoustic survey. Paper SC/50/CAWS20 presented to the
International Whaling Commission, June 1998 (unpublished).
· Best PB (1979) Social organization in sperm whales, Physeter
macrocephalus. pp. 227-289 in H.E. Winn and B.L. Olla (eds.),
Behaviour of marine animals, Vol. 3. Plenum, New York.
· Best PB, Canham PAS, MacLeod N (1984) Patterns of reproduction
in sperm whales, Physeter macrocepahlus. Rep. Int. Whal.
Comm Spec Iss 6:51-79.
· Biggs PB, Jochens AE, Howard NK, Di Marco SF, Mullin KD,
Leben RF, Muller-Karger FE, Hu C (2005) Eddy forced variations in
on- and off-Margin summertime circulation along the 1000-m isobath
of the northern Gulf of Mexico, 2000-2003, and links with sperm
whale distributions around the middle slope. Circulation in the
Gulf of Mexico: Observations and Models. Geophys Mon Ser 161: 71-85.
· Bouquegneau JM, Debacker V, Govert S, Nellissen JP (1997)
Toxicological investigations on four sperm whales stranded on the
Belgian coast: inorganic contaminants. Bul Inst Roy Sci Nat Belgique,
Biologie 67-Suppl.:75-78.
· Carretta JV, Forney KA, Lowry MS, Barlow J, Baker J, Johnston
D, Hanson, Muto MM, Lynch D, Carswell L (2009) U.S. Pacific Marine
Mammal Stock Assessments: 2008. NOAA-TM-NMFS-SWFSC-434
· CETAP. 1982. A characterization of marine mammals and turtles
in the mid- and North-Atlantic areas of the US Outer Continental
Shelf, Final Report. Bureau of Land Management, Washington, DC.
(Ref. AA551-CT8-48).
· Christal J, Whitehead H (1997) Aggregations of mature male
sperm whales on the Galapagos Islands breeding ground. Mar Mamm
Sci 13:59-69.
· Christensen I, Haug T, Øien N (1992) Seasonal distribution,
exploitation and present abundance of stocks of large baleen whales
(Mysticeti) and sperm whales (Physeter macrocephalus) in
Norwegian and adjacent waters. ICES J Mar Sci 49:341-355
· Clapham PJ, Berggren P, Childerhouse S, Friday NA, Kasuya
T, Kell L, Kock K-H, Manzanilla-Naim S, Notarbartolo di Sciara G,
Perrin WF, Read AJ, Reeves RR, Rogan E, Rojas-Bracho L, Smith TD,
Stachowitsch M, Taylor BL, Thiele D, Wade PR, Brownell JrRL (2003)
Whaling as science. Bioscience 53: 210-211.
· Clarke MR (1976) Observation on sperm whale diving. J Mar
Biol Assoc UK 56: 809-810
· Clarke MR, Martins HR, Pascoe P (1993) The diet of sperm
whales (Physeter macrocephalus Linnaeus 1758) off the Azores.
Phil Trans R Soc Lond B 339: 67-82
· Davis RW, Evans WE, Würsig B (2000) Cetaceans, sea
turtles and seabirds in the Northern Gulf of Mexico: distribution,
abundance and habitat associations. OCS Study MMS 2000-03, US Dept
of the Interior, Geological Survey, Biological Resources Division
and Minerals Management Service, Gulf of Mexico OCS Region, New
Orleans, LA.
· Davis RW, Fargion GS, May N, Leming TD, Baumgartner, Evans
WE, Hansen LJ, Mullin AK (1998) Physical habitat of cetaceans along
the continental slope in the northcentral and western Gulf of Mexico.
Mar Mamm Sci 14:490-507.
· Drouot V (2003) Ecology of sperm whales (Physeter macrocephalus)
in the Mediterranean Sea. PhD Thesis, University of Wales, Bangor.
· Engelhaupt DT (2004) Phylogeography, kinship and molecular
ecology of sperm whales (Physeter macrocephalus). PhD Thesis,
University of Durham.
· Engelhaupt D, Hoelzel AR, Nicholson C, Fratzis A, Mesnick
S, Gero S, Whitehead H, Rendell L, Miller P, De Stefanis R, Canadas
A, Airoldi S, Mignucci-Giannoni AA (2009) Female philopatry in coastal
basins and male dispersion across the North Atlantic in a highly
mobile marine species, the sperm whale (Physeter macrocephalus).
Molecular Ecology 18: 4193-4205
· Evans K, Thresher R, Warneke RM, Bradshaw CJA, Pook M,
Thiele D, Hindell MA (2005) Periodic variability in cetacean strandings:
links to large-scale climate events. Biol Lett 1: 147-150
· Forney KA (2007) Preliminary estimates of cetacean abundance
along the U.S. west coast and within four National Marine Sanctuaries
during 2005. U.S. Department of Commerce, NOAA Technical Memorandum
NMFS-SWFSC-406. 27p.
· Fossi MC, Marsili L, Giovanni N, Natoli A, Politi E, Panigada
S (2003) The use of a non-lethal tool for evaluating toxicological
hazard of organochlorine contaminants in Mediterranean cetaceans:
New data 10 years after the first paper published in MPB. Mar Pollut
Bull 46, 972-982.
· Gonzalez E (2001) Interactions of the Dissostichus eleginoides
fishery with Physeter macrocephalus in the central-south
waters of Chile. Abstract in Proceedings of the XXI Marine Sciences
Meeting. Vina del Mar, Chile.
· Goold JC, Whitehead H, Reid RJ (2002) North Atlantic sperm
whale, Physeter macrocephalus, strandings on the coastlines
of the British Isles and Eastern Canada. Can Field Nat 116: 371-388.
· Gordon J, Leaper R, Hartley FG, Chappell O (1992) Effects
of whale-watching vessels on the surface and underwater acoustic
behaviour of sperm whales off Kaikoura, New Zealand. Science &
Research Series No. 52, Department of Conservation, Wellington,
N.Z. 64 pp.
· Gunnlaugsson T, Sigurjónsson J (1990) NASS-87 :
Estimation of whale abundance based on observations made onboard
Icelandic and Faroese survey vessels. Rep Int Whaling Commn 40:
571-580
· Ivashin MV, Rovnin AA (1967) Some results of the Soviet
whale marking in the waters of the North Pacific. Norsk Hvalf-Tid
56:123-135.
· Jacques TG, Lambertsen RH (eds.) (1997) Sperm whale deaths
in the North Sea: science and management. Bull Ins Roy Sci Nat Belgique,
Biologie 67-Suppl.: 133pp.
· Jaquet N (1996) How spatial and temporal scales influence
understanding of sperm whale distribution: a review. Mamm Rev 26:51-65.
· Jaquet N, Dawson S, Slooten E (1998) Diving behaviour of
male sperm whales: foraging implications. International Whaling
Commission, Scientific Committee Doc. SC/50/CAWS 38, 20 pp.
· Jaquet N, Whitehead H (1996) Scale-dependent correlation
of sperm whale distribution with environmental features and productivity
in the South Pacific. Mar Ecol Prog Ser 135:1-9.
· Jefferson TA, Webber MA Pitman RL (2008) Marine mammals
of the world. Elsevier, Amsterdam, 573 pp.
· Jensen AS, Silber GK (2004) Large whale ship strike database.
NOAA Tech Memo NMFSOPR-39 pp.
· Jochens A, Biggs D, Benoit-Bird K, Engelhaupt D, Gordon
J, Hu C, Jaquet N, Johnson M, Leben R, Mate B, Miller P, Ortega-Ortiz
J, Thode A, Tyack P, Würsig B (2008) Sperm whale seismic study
in the Gulf of Mexico: Synthesis report. U.S. Dept. of the Interior,
Minerals Management Service, Gulf of Mexico OCS Region, New Orleans,
LA. OCS Study MMS 2008-006. 341 pp.
· Johnson M, Tyack P, Miller P, Nowacek D (2003) DTAG: A
synchronous sound and orientation recording tag for wild marine
mammals. Environmental Consequences of Underwater Sound (ECOUS)
Symposium. 12-16 May 2003. San Antonio, Texas.
· Kanwisher JW, Ridgway SH (1983) The physiological ecology
of whales and porpoises. Sci Am 248: l10-120
· Karpouzli E, Leaper R (2004) Opportunistic observations
between sperm whales and deep-water trawlers based on sightings
from fisheries observers in the northwest Atlantic. Aquat Cons:
Mar & Freshw Ecos 14: 95-103.
· Kasuya T, Miyashita T (1988) Distribution of sperm whale
stocks in the North Pacific. Sci Rep Whales Res Inst 39: 31-75.
· Kato H, Miyashita T (1998) Current status of North Pacific
sperm whales and its preliminary abundance estimates. Unpubl. report
submitted to Int. Whal Commn (SC/50/CAWS/52). 6 pp.
· Kato H, Miyashita T (2000) Current status of the North
Pacific sperm whale and its preliminary abundance estimates. Paper
SC/50/CAWS2 presented to the Scientific Committee of the Int. Whal
Commn
· Kawakami T (1980) A review of sperm whale food. Sci Rep
Whales Res Inst Tokyo 32: 199-218
· Laist DW, Knowlton AR, Mead JG, Collet AS, Podesta M (2001)
Collisions between ships and whales. Mar Mamm Sci 17: 35-75.
· Lambertsen RH (1990) Disease biomarkers in large whales
of the North Atlantic and other oceans. Pp. 395-417 in J.F. McCarthy
and L.R. Shugart (eds.), Biomarkers of environmental contamination.
Lewis Publishers, CRC Press, Boca Raton, FL.
· Law RJ, Morris RJ, Allchin CR, Jones BR (1997) Metals and
chlorobiphenyls in tissues of sperm whales (Physeter macrocephalus)
and other cetacean species exploiting similar diets. Bull Instit
Roy Sci Nat Belgique, Biologie 67-Suppl.:79-89.
· Lyrholm T, Leimar O, Johanneson B, Gyllensten U (1999)
Sex biased dispersal in sperm whales: contrasting mitochondrial
and nuclear genetic structure of global populations. Proc Roy Soc
Lond B 266: 347-354.
· Madsen PT, Payne R, Kristiansen NU, Wahlberg M, Kerr I,
Møhl B (2002) Sperm whale sound production studied with ultrasound
time/depth-recording tags. J Exp Biol 205:1899-1906.
· McCall HMP (1999) Sperm whales, Physeter macrocephalus,
in the Gully, Nova Scotia: population, distribution, and response
to seismic surveying. BSc Hons Thesis, Dalhousie University, Halifax,
Nova Scotia.
· Mitchell E (1975) Preliminary report on Nova Scotia fishery
for sperm whales (Physeter catodon). Rep Int Whal Commn 25:
226-235.
· Mullin K, Hoggard W, Roden C, Lohoefener R, Rogers C, Taggart
B (1991) Cetaceans on the upper continental slope in the north-central
Gulf of Mexico. OCS Study/MMS 910027. U.S. Department of the Interior,
Minerals Management Service, Gulf of Mexico OCS Regional Office,
New Orleans, Louisiana. 108 pp.
· Mullin KD (2007) Abundance of cetaceans in the oceanic
Gulf of Mexico based on 2003-2004 ship surveys. 26 pp. NMFS, Southeast
Fisheries Science Center, Pascagoula, USA.
· Mullin KD, Fulling GL (2004) Abundance of cetaceans in
the oceanic northern Gulf of Mexico, 1996-2001. Mar Mam Sci 20:
787-807.
· NMFS (2009) Sperm Whale (Physeter macrocephalus.
5-Year Review: Summary and Evaluation. National Marine Fisheries
Service Office of Protected Resources, Silver Spring, MD, 44. pp.
· Palacios DM, Gerrodette T (1996) Potential impact of artisanal
gillnet fisheries on small cetacean populations in the eastern tropical
Pacific. NOAA Admin Rep LJ-96-11.
· Purves MG, Agnew DJ, Balguerias E, Moreno CA, Watkins B
(2004) Killer whale (Orcinus orca) and sperm whale (Physeter
macrocephalus) interactions with longline vessels inChe Patagonia
toothfish fishery at South Georgia, South Atlantic. CCAMLR Science
11 (111-126).
· Reeves RR (2002) The origins and character of 'aboriginal
subsistence' whaling: a global review. Mam Rev 32: 71-106.
· Reeves RR, Notarbartolo Di Sciara G (2006) The status and
distribution of cetaceans in the Black Sea and Mediterranean Sea.
IUCN Centre for Mediterranean Cooperation, Malaga, Spain.
· Rice DW (1989) Sperm whale Physeter macrocephalus
Linnaeus, 1758. in Ridgway SH, Harrison R (eds.), Handbook of marine
mammals, vol. 4. Academic Press, London. Pp. 177-233
· Richter C, Dawson S, Slooten E (2006) Impacts of commercial
whale watching on male sperm whales at Kaikoura, New Zealand. Mar
Mam Sci 22:46-63.
· Santos MB, Pierce GJ, García Hartmann M, Smeenk
C, Addink MJ, Kuiken T, Reid RJ, Patterson AP, Lordan C, Rogan E,
Mente E (2002) Additional notes on stomach contents of sperm whales
Physeter macrocephalus stranded in the north-east Atlantic. J Mar
Biol Assoc UK 82: 501-507
· Simon MJ, Kristensen TK, Tendal OS, Kinze CC, Tougaard
S (2003) Gonatus fabricii (Mollusca, Theuthida) as an important
food source for sperm whales (Physeter macrocephalus) in
the Northeast Atlantic. Sarsia 88: 244-246.
· Stone CJ (2003) The effect of seismic activity on marine
mammals in UK waters, 1998-2000. Report No. 323 of the Joint Nature
Conservation Committee, Aberdeen, Scotland.
· Taylor BL, Baird R, Barlow J, Dawson SM, Ford J, Mead JG,
Notarbartolo di Sciara G, Wade P, Pitman RL (2008) Physeter macrocephalus.
In: IUCN 2009. IUCN Red List of Threatened Species. Version 2009.2.
<www.iucnredlist.org>.
· Vanselow KH, Ricklefs K (2005) Are solar activity and sperm
whale Physeter macrocephalus strandings around the North
Sea related? J Sea Res 53: 319- 327.
· Vanselow KH, Ricklefs K, Colijn F (2009) Solar driven geomagnetic
anomalies and sperm whale (Physeter macrocephalus) strandings around
the North Sea: an analysis of long term datasets. The Open Mar Biol
J 3: 89-94.
· Viale D, Verneau N, Tison Y (1992) Stomach obstruction
in a sperm whale beached on the Lavezzi islands: Macropollution
in the Mediterranean. J Rech Oceanogr 16: 100-102.
· Wade PR, Gerrodette T (1993) Estimates of cetacean abundance
and distribution in the eastern tropical Pacific. Rep Int Whal Commn
43:477-493.
· Waring GT, Josephson E, Fairfield CP, Maze-Foley K (2005)
U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments
-- 2005. NOAA Tech Memo NMFS-NE-194: 318pp.
· Waring GT, Josephson E, Fairfield-Walsh CP, Maze-Foley
K (eds.) (2007) U.S. Atlantic and Gulf of Mexico Marine Mammal Stock
Assessments -- 2007. NOAA Tech Memo MFS NE 205; 415 p.
· Waring GT, Josephson E, Fairfield-Walsh CP, Maze-Foley
K, eds (2009) U.S. Atlantic and Gulf of Mexico Marine Mammal Stock
Assessments -- 2008. NOAA Tech Memo NMFS NE 210; 440 p.
· Waring GT, Palka DL, Mullin KD, Main JHW, Hansen LJ, Bisack
KD (1997) U.S. Atlantic and Gulf of Mexico marine mammal stock assessments
- 1996. NOAA Tech Memo NMFS-NE-114: 250pp.
· Watwood SL, Miller P, Johnson M, Madsen PT, Tyack PL (2006)
Deep-diving foraging behaviour of sperm whales (Physeter macrocephalus).
Journal of Animal Ecology 75: 814-825.
· Weir CR (2008) Overt responses of humpback whales (Megaptera
novaeangliae), sperm whales (Physeter macrocephalus),
and Atlantic spotted dolphins (Stenella frontalis) to seismic
exploration off Angola. Aquat Mamm 34: 71-83
· Whitehead H (2002) Estimates of the current global population
size and historical trajectory for sperm whales. Mar Ecol Prog Ser
242:295-304.
· Whitehead H (2003) Sperm whales: social evolution in the
ocean. University of Chicago Press, Chicago, IL.
· Whitehead H (2009) Sperm whale Physeter macrocephalus.
In: Encyclopedia of marine mammals 2nd Ed. (Perrin WF, Würsig
B, Thewissen JGM, eds.) Academic Press, Amsterdam, 1091 - 1097.
· Whitehead H, Brennan S, Grover D (1992a) Distribution and
behaviour of male sperm whales on the Scotian Shelf, Canada. Can
J Zool 70: 912-918.
· Whitehead H, Waters S, Lyrholm T (1992b) Population structure
of female and immature sperm whales (Physeter macrocephalus)
off the Galapagos Islands. Can J Fish Aquat Sci 49:78-84.
· Wright A (2005) Lunar cycles and sperm whales (Physeter
macrocephalus) strandings on the North Atlantic coastlines of
the British isles and eastern Canada. Mar Mam Sci 21: 145-149.
© Boris Culik (2010) Odontocetes.
The toothed whales: "Physeter macrocephalus". UNEP/CMS
Secretariat, Bonn, Germany. http://www.cms.int/reports/small_cetaceans/index.htm
© Illustrations by Maurizio Würtz, Artescienza.
© Maps by IUCN.
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