Review
of the great white shark Prepared by Anthony
Gibbins Conservation:
Great white shark Class Chondrichthyes Common names: White shark; maneater
(USA), white pointer, white death (Aus.), blue pointer, Tommy shark,
Uptail (S. Africa). French: Grand requin blanc; requin blanc; Lamie
(Nice) Current IUCN Status Assessment: VULNERABLE globally • Taxonomy & Early History Only one white shark species recognised by taxonomists - but it was not
always known as Carcharodon carcharias, the 'Jagged-Toothed One'.
Carolus Linnaeus, in his 1758 tome Systema Naturae, named it
Squalus carcharias and thereafter the species was afforded a
variety of binomials, including Carcharias lamia, Carcharias verus,
Carcharodon smithii and Carcharodon rondeletii. This shark was known from the Mediterranean in ancient times, most
probably being the fish referred to by Aristotle and other Greek writers
as the fearsome Lamia monster - a common-name still used for this
species both in Greece and in places along the coast of southern France
(where it is called the 'Lamie'). They were not uncommonly caught between
Sète and Nice in medieaval centuries, and specimens were of as much
interest to the writers of the time as they are today. In 1566, the
Montpellier-based naturalist Guillaume Rondelet noted the voracious
appetite of the Lamia ; a diet which, he commented, included tuna
and even the odd human - and suggested it was this animal, rather than a
whale, that was responsible for swallowing the prophet Jonah in the famous
biblical fable. It's not hard to see where Rondelet's reasoning
originated. The white shark was then - as it is now - a little-known,
rarely seen but greatly feared animal of mythical status, whose apparent
penchant for consuming humans (in a mystical sense!) was well-known
amongst the region's seafarers. It seems quite conceivable that, on some
ancient shoreline and unrecorded date, a white shark was caught and
eviscerated, leading to the rare discovery of human remains, perhaps
largely intact and fresh, in its stomach (e.g., see Conorelli &
Perrando, 1909, for a genuine example of this). Such an event would
unquetionably spark considerable public interest and quickly evolve into
local fable. Perhaps inserting the 'whale' element gave the story a more
benign feel, as befitting the religious context into which it is now
told. Throughout much of the 18th and 19th Century, there was confusion in
the proper identity of the white shark amongst the Mediterranean
elasmobranch fauna. Its triangular, serrated teeth proved a cornerstone in
fuelling the nomenclaturial and descriptive mayhem that followed - not
least as a much more common Mediterranean species, the sandbar shark
Carcharhinus plumbeus, has teeth that may - at least when compared
solely by cursory, written description - fit adequately into the white
shark 'mould'. The thought of the essentially inoffensive sandbar shark
being unwittingly catapulted to 'maneater' status may seem amusing in
hindsight but, at the time, such fundamental errors of identification
generated many misrepresentative species accounts. The white shark was
often grouped collectively with carcharhinids on account of their rather
similar dentition. Poor illustrations of the white shark also abounded,
exasperating the problem. The simple fact was that very few scientists
describing this animal had actually seen one, either dead or alive, and
much of the topical knowledge explicating both its physical appearance and
behaviour drew more upon hearsay and mariner's lore than biological
fact. Marcus Bloch's (1785-95) illustration of Squalus carcharias is a
typical case-in-point. The excised jaws are depicted upside-down - a
fundamental mistake later to be followed blindly by other authors - and
the shark itself bears only a limited resemblance to the fish we call
Carcharodon carcharias. With an asymmetrical tail, small gill-slits
and sail-like first dorsal fin, it might come as no surprise that the
drawing has more than a passing similarity to Carcharhinus plumbeus.
Arguably, it is only the single tooth, illustrated alongside, that can
be readily attributed to the white shark. Meanwhile, Bloch's white shark
"mugshot" doubtlessly condemned yet more benign sandbars to the maneater's
Hall of Fame. It is poignant to add that a much earlier depiction,
prepared by Guillaume Rondelet in 1554, was far more accurate than many
later images - correctly showing a crescentic caudal fin and caudal keels.
Even the relative positions of second dorsal and anal fins are
precise. Similarly, C.L. Bonaparte (1832-1841) illustrated the white shark in
his description of Italian vertebrate fauna. His rendition is essentially
accurate and almost certainly based upon a freshly-caught specimen.
Interestingly, the black axillar blotch so often found on these sharks is
missing from the picture - and indeed is often absent on many
Mediterranean examples. In 1838 L. Agassiz, a paleontologist, published a description of the
genus now called Carcharodon with some precision in a catalogue of
fossil fish, naming the living species as Carcharodon smithii. He
had adopted the new generic name following the British naturalist and
physician Dr. Andrew Smith who had, at some time whilst resident in South
Africa between 1820 and 1836, procured a small white shark specimen from
the Cape Province region. Smith was the first scientist to properly
distinguish the genus and proposed the generic name Carcharodon in
an 1838 work by the German anatomists, Johannes Müller and Fredrich Henle.
In an 1839 discussion of white sharks, Müller and Henle again followed
Smith's generic name and called the fish Carcharodon rondeleti.
They suggested that the South African shark was somewhat different
from other white sharks and might thus represent a second species. As it happened, Andrew Smith was to personally describe his specimen in
a later publication of 1849, naming it as Carcharodon capensis in
reference to its capture-locality. He mentioned stomach-contents and noted
aspects of white shark behaviour with an air of reality far divorced from
the mediaeval-inspired writings and mythos that existed before. In 1851,
J.E. Gray synonymised Smith's animal with C. rondeleti, realising
that this was one and the same species, rather than a unique South African
endemic. This assertion has been followed ever since. For decades, Smith's stuffed C. capensis holotype - a juvenile
female measuring just over 2 metres in length - lay gathering dust and
more than a few errant white paint-splashes in the storage basement hall
of the British Museum of Natural History, London - until re-discovered
there in June 1994 by Leonard Compagno, Oliver Crimmen (of the museum's
Fish Section) and the author. Manoeuvring the hefty specimen down from
it's ceiling-high resting place on a top shelf, and then lifting it up
two flights of stairs to a laboratory, was an exhausting episode taking
perhaps more physical effort than the process of actually catching it
in the first place. Despite showing its age, the shark still had a legible
pencilled label affixed to its wooden base, reading "Carcharodon capensis,
Cape Seas". Preservation was sufficiently good to even see the black
axillar blotch beneath each pectoral fin insertion. The anal fin, however,
was missing - perhaps accidentally destroyed during the mounting process. • Status and Biological
Summary A fusiform, heavy-bodied shark with a crescentic caudal fin and large,
triangular, coarsely serrated teeth. In juveniles under 1.8 m, the teeth
have small lateral cusplets and in neonates, anterior lower-jaw teeth may
essentially lack marginal serrations. Typical tooth-count is:- The snout is conical, blunt and dorsally flattened; the five gill-slits
are large but not encircling the head. The first dorsal fin, nearly an
equilateral triangle in shape with a slightly concave rear margin, has its
origin opposite, or slightly anterior, to the inner corner of the pectoral
fins. The first dorsal is rounded at the apex in neonates but becomes more
acutely-pointed within the first two years. The second dorsal fin is
minute and pivotable, its posterior margin above or slightly ahead of the
origin of the equally diminutive anal fin. The broad, dorsoventrally
depressed caudal peduncle is expanded laterally to form a prominent keel
on either side, but without secondary keels on the ventral caudal lobe
itself (as in the porbeagle Lamna nasus). The lunate caudal fin tends to
have acutely-pointed tips in sharks over 2.0 m TL. In neonates, the lower
(ventral) caudal surfaces may be more rounded and compressed but rapidly
expand soon after birth. The caudal lobes are almost of equal size, with
the lower (ventral) anterior margin measuring 76-92% of the same
measurement on the upper (dorsal) lobe. The pectoral fins are large and
moderately falcate. • Colour Size and
Reproduction Lengths at maturity for both sexes remain somewhat undetermined, and
based on (currently limited) age-growth data it may be possible that
different populations mature at varying lengths. The majority of females
mature at between 450-500cm TL (Francis, 1996), but have been reported as
immature at sizes as much as 472cm (Springer, 1939). Males mature at about
350-360cm (Pratt, 1996). A single study of age and growth, pooled from 21
specimens (Cailliet et al., 1985) suggests a generalised age of maturity
of 10-12 years. A mature female of 500cm is estimated to have reached ca.
14 to 16 years. • Weight-at-Length
relationship In W= 2.0686 + 3.0958 In TL (W in kg, TL in m) y on x regression: In W= 2.1154 + 3.0542 In TL (N = 327; r2 =
0.973) A further WT-TL regression was given by Compagno (1984), based on 98
specimens (mainly from California, with a range of 127 to 554cm): WT = 4.34 x 10-6 TL 3.14 •
Reproduction Verified litter-sizes have ranged from 5 to 10
near-term foetuses, with unconfirmed reports of up to 14 embryos. Size at
birth, based on examined foetuses and free-swimming neonates, is within a
range of 120-150cm TL. The reproductive mode is viviparity without a
placenta and embryos are nourished by oophagy - the ingestion of
unfertilised eggs. Whilst in-utero, the embryonic white sharks swallow
their own sets of shedded teeth, perhaps to re-utilise calcium and other
minerals. Gestation times are unknown but doubtless long - close to a
year, perhaps. It is possible that any one female only reproduces
biennially, mating soon after giving birth, but this remains to be
confirmed. Some incidents of fresh and healed bite-marks found
on the dorsum, flanks and particularly the pectoral fins of mature female
white sharks has been interpreted as an indicator of mating activity,
wherein males have grasped the female during copulation (as witnessed with
other species of sharks). Copulation has not been reliably witnessed in
this species, although an observation was reported by a volunteer seal
researcher describing the curious surface behaviour seen between two great
whites off the Nuggets, on the Otago Peninsula of New Zealand's South
Island, that may have been mating (Francis, op. cit.). Scrutiny of worldwide records of pregnant females,
coupled with the localised, seasonal abundance of newborns, suggests that
parturition occurs during the Spring to late Summer in temperate shelf
waters. Areas where large adults of both sexes and neonates occur together,
indicating their likely significance for reproduction and as nursery-grounds,
include the shelf waters off the Northeastern United States (the Mid-Atlantic
Bight); Southern California and Baha; Southern and Eastern Australia;
New Zealand; Japan; Eastern Cape Province of South Africa and the Southern-Central
Mediterranean Sea, especially the waters between Western Sicily and Tunisia.
Young white sharks of a year-old or less have also been caught elsewhere
in the Mediterranean - off Algeria, France and in the North Aegean, for
example. The majority, however, originate from the Sicilian Channel during
high summer, when Sicilian-based trawlers net the young sharks on the
seabed at surprising depths of up to about 200 m. White sharks occupy a cosmopolitan distribution throughout temperate
seas and oceans and will occasionally penetrate tropical zones. Sporadically,
they make incursions to cold, boreal waters and have been recorded from
off the south Alaskan and Canadian coasts (Gulf of St. Lawrence). Principally
an epipelagic dweller in neritic waters, the white shark is found from
the surfline to well offshore, often in association with isolated islets
and archipelagos inhabited by pinnipeds (e.g., the Farallon Islands, Marin
Co., California; Neptune Islands and Dangerous Reef, Spencer Gulf, South
Australia; Dyer, Seal and Bird Islands, South Africa), rocky headlands
where deepwater lies close to shore (California and Oregon coasts; Central
Chile; Ligurian Sea, Italy), or near offshore fish reefs and shoals (e.g.,
Adventure Bank, Sicilian Channel; Struis Bay area, Cape Agulhas, South
Africa). A synopsis of confirmed worldwide occurrences are listed below by
geographic region. Indices of relative abundance (based on
capture-records, sightings, interactions with man) are abbreviated thus:
CA: Center of Abundance F: Frequent; O: Occasional; I: Infrequent; R:
Rare; VR: Very Rare. Further remarks are added where relevant. Northwest Atlantic and Caribbean:- Gulf of St Lawrence,
Newfoundland, St. Pierre Bank, Sable Island, Bay of Fundy (O-R); New
Brunswick, Maine, Massachusetts, Rhode Island, Martha's Vineyard, Long
Island, New York coastline and offshore, particularly within the 30-fathom
contour (CA; F); south along US coast through the Carolinas to Florida
(R-O). Gulf of Mexico, principally western Florida shores (I) but ranging
past the Mississippi Delta to at least Corpus Christie, Texas (R);
Caribbean occurrences - Florida Keys, Cuba (Cojimar), Bahamas
(R). Southwest Atlantic:- Central and Southern Brazil (R-VR), 10
examples known to 1993 of which 6 were from Rio de Janeiro State, 3 from
Ceará State and 1 from Espírito Santo State; Argentinean coast at Puerto
Quequén (Necochea) and south to at least C. Blanco (R). Northeast Atlantic:- Charente-Maritime coast of France (La
Rochelle and Pertuis d'Aintoche; VR), north to the mouth of the Loire (VR,
1 old record); Gulf of Gascogne, Northern Spanish and Portuguese coast
southwards to Tarifa and Gibraltar (VR). Azores Archipelago (I); Madeira
(VR). Morocco, Mauritania, to Senegambia (Dakar region at Île de Gorée and
environs), Ghana and Cape Verde Is. (R). Canary Islands at Gran Canaria
and Tenerife (VR?); probably Lanzarote. Possibly Zaire but literature
confusion likely with bull sharks, Carcharhinus
leucas. Mediterranean Sea (CA): Spain and Balearics, including Catalon
Sea at Valencia, Vinaroz and Islas Columbretes; also recently (1992) from
Tossa de Mar and Andraitx, Majorca; Cabo Salinas, Majorca; Menorca (R).
Gulf of Lyons, particularly Sète, Palavas and Grau-du-Rois (I-O); Riviera
and Côte D'Azur (I), Ligurian Sea, especially from Camogli and Portofino
to La Spezia (O-R). Western Italy (Tyrrhenian Sea: Elba, Anzio, San Felice
Circeo, Ponza, Lipari Islands, Naples, Calabria Region), Corsica, Sardinia,
especially northern part at Capo Testa; southwest region at Isola di San
Pietro (O); Morocco and Melilla (O); Algerian coast (I-O?), Tunisia, principally
La Galite Islands, Cape Bon area and Gulf of Gabès near Kerkennah Islands
and Djerba (O), Sicily (cosmopolitan, especially Egadi Islands, Trapani
Region and Messina); Sicilian Channel, Pantelleria, Isole Pelagie and
offshore banks (F-O); Malta and Gozo (I). Ionian Sea off Calabria (Capo
Spartivento) and Galipolli. Southern and Central Adriatic off Puglia,
Manfredonia, Ancona and Termoli; also Dalmatian coast near Split (I-O),
Northern Adriatic, principally Istria and Kvarner Gulf of Croatia (I,
formerly F), sporadically at Riccione, Rimini and in the Gulfs of Venice
and Trieste. Corfu, Greek Mainland, Aegean Sea (Thermaikos Gulf, Thassos,
Kavalla, Alexandroupolis) through Turkish coast (Foça region) to Bosphorus
(R) but not Black Sea. Cyprus (O), Israel ('Akko) and Egypt (Agamy, near
Alexandria) (R). Southeast Atlantic:- Gough Island (single recent record).
Namibia, from Cape Cross and Swakopmund southwards (O, but little data)
possibly north to mid-Angola?, entire Cape Province, especially eastwards
of the Cape of Good Hope in False Bay, Walker Bay, Gansbaai, Danger Point,
Kleinbaai, Dyer Island group (CA; F), Cape Agulhas region (Struisbaai, Die
Mond, Arniston) and entire Wild Coast (F) to Port Elizabeth; Algoa Bay
(especially Bird Island) to Natal (F); northerly extent of range along the
Indian Ocean coast of Africa uncertain. Indian Ocean:- Northern Natal coast (F) but more sporadic
towards tropical extremes, perhaps more extensive around Southern
Mozambique and Madagascar than previously supposed, based on recent
sightings. Kenyan coast at Malindi; limits to northerly range in Western
Indian Ocean currently unclear. Appears absent in the Red Sea (one old,
doubtful record) or a very occasional outlier at best. Kuwait coast and
Persian Gulf (VR); Seychelles Island (VR), Cargados archipelago (VR); Sri
Lanka (VR). Indo-Pacific and Western Pacific:- Kamchatka region (VR,
possibly dubious), Japanese Islands (Honsu and Hokkaido, including Inland
Sea) (CA; O); Okinawa (I), Eastern Chinese Coast (VR), Korean Peninsula at
Pusan (VR), Taiwan at Keelung (VR); possibly South China Sea along the
Chinese coast and off Hong Kong (VR, dubious); Coral Sea (VR); Philippines
(Palawan, Mindanao) and Bonin Island (VR). Australia: Queensland (F-O),
entire coastline to NSW and South Australia (CA; F), Tasmania (F-O), Great
Australian Bight (CA; F), Western Australia (CA; F-O). New Zealand, both
North and South Islands (CA; O), particularly Dunedin, Otago peninsula and
environs on South Island (F-O). Stewart Island, Chatham Islands (O);
Campbell Island, recent attack there upon a diver (I-R?). New Caledonia
(VR). Central Pacific:- Hawaiian Islands off Oahu, Hawaii and
Northwestern Islands (R); Marshall Islands at Bikini (VR); Easter Island,
based on recent shark attack. Northeast Pacific:- Gulf of Alaska (VR), British Columbia,
Washington State (R), Oregon Coasts (O), abundance increases southwards to
Northern California (CA; F), Central California (CA; F, particularly from
Sonoma County, Marin Co., the Gulf of the Farallones, Año Nuevo Island,
Santa Cruz, Monterey Peninsula to the Big Sur); Southern California and
Channel Islands (CA; F-O), Baha and offshore at Isla de Guadalupe (O). Sea
of Cortez (I-R), Mexico (R) to Panama (R). Southeast Pacific:- From Panama south through Ecuador and Peru
to Chile; data sketchy, however, for range north of Chile. Northern and
Central Chile (CA; O) to Archipelago de los Chonos (45úS), possibly more
southerly to Cape Horn. • Abundance • Movements and
Nomadicy Tracking of white sharks by telemetry has afforded an insight into facets
of their spatial behaviour, both in the horizontal and vertical sense.
An efficient, graceful cruiser at slow speeds (averaging 3.2km/h over
a distance of 190km, in a classic experiment off New York by the late
Frank Carey et al., 1982), the white shark, in keeping with other lamnids,
maintains elevated temperatures within the swimming muscles, brain, eyes
and gut to at least 13.7ûC above the surrounding sea temperature - an
adaption which may provide an increase in neural, digestive and muscle
activity and function. The species is also capable of short-duration,
high-velocity pursuits, even launching entirely clear from the surface
in spectacular repeated leaps. 'Blind' traditional telemetry and more
recent remote videography using parasitic 'Crittercam' video cameras reveals
that white shark predominantly cruises in a purposeful manner, either
just off the bottom or near to the surface, but spending very little time
at midwater depths unless stimulated by baits (Strong et al., 1992; Goldman
et al., 1996; Greg Marshall, pers. comm.). This species is freely capable of making long-distance movements on localised,
regional and intercontinental scales. Although information is rather limited
by the essential rarity of these animals, some data on movements has been
afforded through tag-and-release programmes in America, South Africa and
Australia. For example, two white sharks tagged by a joint Cousteau Society
and South Australian Department of Fisheries (SADF) Team near Dangerous
Reef were later recaptured some 220km to the east, near Antechamber Bay
on Kangaroo Island. Similarly, a South African specimen was found to have
travelled approximately 780km in just 28 days. But whilst tagging has
demonstrated these rather long-distance movements, the same studies have
also highlighted an interesting propensity for white sharks to exhibit
a high degree of localised, temporary residency. At the Farallon Islands,
a stark archipelago some 40km due west of San Francisco, researchers have
identified a number of individual adult white sharks through distinctive
body-markings and fin abnormalities - rather as accomplished in equatable
studies of whales and dolphins. Over the course of successive yearly field
studies on the island since the early 1970's, it has been shown quite
conclusively that some of the Farallon sharks return to the locale annually
in the Autumnal months, often arriving and departing on virtually the
same repeated days or weeks with surprising homogeneity. White sharks
are scarce at the islands between January and September, but precisely
where those seen in Autumn move away to through the remainder of the year
is presently unknown. Comparable recent work at Dyer Island, South Africa
and Dangerous Reef has yielded similar results to the Californian study
- some tagged (and thus readily identifiable) sharks returning repeatedly
to the same places, where they are then resighted by observers with considerable
frequency. Nevertheless, an even greater proportion of tagged sharks are
never resighted again - testimony to the nomadic side of these fish. Predatory Biology Social
and Complex behaviour
Threats • Directed Fisheries • Indirect Mortality
and Beach Meshing Somewhat more directed pressure is manifested in the form of anti-shark
beach netting, installed to ensure the safety of bathers at selected localities.
Shark nets have been installed at some Australian beach resorts in New
South Wales since 1937, and along Queensland's Gold Coast since 1962.
Similar measures are taken in South Africa where in 1952, responding to
a series of high-profile local shark attacks, the first nets were installed
off Durban. This was followed by the installation of two further nets
at Amanzimtoti in August 1962. As of 1990, 14% of the 326 km of Natal
coastline between Mzamba and Richard's Bay was protected by 44.4 km of
gillnets (Dudley and Cliff, 1993), with the operation controlled by the
Natal Sharks Board. In the Queensland Shark Control Programme and NSW meshing operation,
there is data to indicate an irregular decline in white shark captures
over the past three decades. The Natal net-catches - a total of 591 white
sharks between 1974 and 1988, with from 22 to 61 examples caught annually
(Cliff et al., 1989) - indicates a similar but less clearly-defined downward
trend. Conscious of mounting environmental concern and criticism, there
is an increasing willingness by meshing contractors to tag and release
sharks found alive in the nets, although mortality remains high overall
both to sharks and other inoffensive marine creatures including rays,
guitarfish, dolphins and turtles. The broad effect of these meshing programmes
on marine ecology - not least because they are totally unselective by
its very nature - has created considerable controversy and some current
research is directed at developing harmless alternative measures, such
as electronic barriers to repel sharks. • Habitat Loss or Degradation
•Other
Factors Originally, cage diving and white shark viewing blossomed in Australia,
following the success of early endeavours by filmmakers of the 1960's and
1970's including Ron and Valeria Taylor, Stan Waterman, Peter Gimbel and
others (e.g., Blue Water, White Death). South African efforts came
later through operations in False Bay and elsewhere on the Cape Coast, but
are arguably are now more prominent given that reported successes of
Australian 'pay-for-view' operations seemed to diminish in the 1990's, as
indeed did the Spencer Gulf's population of white sharks. In the 1990's,
Californian cage diving operations started opportunistically near Santa
Cruz but were banned by legislation in the wake of fierce local opposition
by surfers groups, scientists and others. All deliberate attraction of
white sharks in Californian waters has since been outlawed by state
legislation, and no cage diving operation directed solely at white sharks
now exists there. In South Africa, a variety of commercial cage tour operators have jostled
for position and clientele - sometimes quite literally so - in the narrow
confines of the waters that separate Dyer Island from Geyser Rock. In
recent months, local conservation authorities and regulatory bodies expressed
concerns over the increasingly 'circus' atmosphere prevailing at the locale,
sometimes coupled with operator's indifference to the area's status as
a nature reserve, and the process has resulted in a temporary moratorium
on various white shark related activities there.Various activities that
could ultimately prove injurious to sharks, such as repeated biopsy sampling,
have been included in the temporary ban and all further applications to
pursue research at the islands must be first scrutineered by local recognised
experts (Chondrichthyan Working Group). The motivation behind the current
moratorium is perhaps best explained by Leonard Compagno (pers.comm),
who draws parallels to the protection of South Africa's terrestrial predators.
"If I were to pull-up in my car at Kruger Park," says Compagno, "and start
firing dart tags and such like at lions or cheetahs, without any sort
of research permit to so so, I'd be quickly arrested." In other words,
the current closely-guarded and vetted access to lions, cheetahs and other
terrestrial mammalian predators should be quite properly extended to an
equally protected marine piscine predator, the white shark. Few would
disagree with this rationale. Whilst cage diving can clearly benefit the white shark in many ways,
and offer a more passive and profitable incentive to former game or commercial
fishermen, the South African example demonstrates that as much attention
should be given to vetting and licensing such activities as is afforded
to banning directed fishing itself. • Present and Planned action In April 1996, a status report on the great white shark was prepared
for the World Conservation Union's (IUCN) 'Red List' - a published
compendium that details threatened animals and plants, offering the sort
of definitive information that may aid in the planning of conservation
strategies. That assessment (see Part 3) concluded that the fish should be
enlisted then under the banner of 'Vulnerable', but that given more data,
this category may change to "Engangered' on at least a regional basis. The
report also concluded that only a global solution will ultimately
safeguard this very global fish. Nevertheless, this is not to discourage such conservation efforts - far
from it. Unilateral or regional attempts to conserve great white sharks
are laudable, as they may provide the encouragement for neighbours to
adopt commensurate measures. The situation can change rather rapidly,
as was the case in Australia. At the time of writing, the Shark Trust
and EEA were pressing the Maltese Government to protect white sharks in
their coastal waters, and become the first Mediterranean nation to take
such an inportant step in the process. Such a high-profile move may even
help advance the long-delayed ratification of the Barcelona Convention,
which lists the white shark in it's appendices as an endangered fish,
needing urgent protection. The pitfalls of inadequate enforcement must be emphasised. Analogous to
the unlawful poaching of rhinos or tigers for traditional medicines, the
demand for procuring white shark carcasses can only be nullified through
equally stringent efforts to banish the market for their end-product
curios. Only pan-national legislation will provide this long-term
security, such as enacted through the jurisdiction of CITES enlistment. In
September 1998, representatives from the Humane Society indicated that
Australia would pursue a CITES enlistment for the white shark, a move
since confirmed and which deserves wider international support -
especially so from both South African and the USA. Given the traditionally
negative status of great white sharks within the public psyche, any
public-supported protection of this enigmatic nimrod is a ray of hope in
aspiring to conserve less well-known, less fearsome - but no less equally
threatened - elasmobranch species. Proposed International CITES Listing: The way
forward? The white shark story is one that charts the very worst of human ignorance,
indifference, insecurity and greed, where it destroys and neglects our
wildlife and environment. This is not a fish to be underestimated. It
is neither a bloodthirsty, mindless killer nor a playful, good-natured
puppy-dog. It is a sentient, intelligent predator deserving of our rational
respect like any other potentially lethal animal. For confirmation of
this, ask any fur seal... Just a couple of quick Pictures to celebrate the greatest suspense film
ever.. Meet 'BRUCE', the 7.3 meter mechanical shark which was built for the
film Jaws. It weighs 1.5 tonnes and cost about £100,000 to make,
in all 3 were made. Examining the model is world renowned shark expert
Valerie Taylor.
Carcharodon carcharias
Contents
Status
& Biology:
• Nomenclature & Taxonomy
• Description
• Size
• Weight-Length relationship
• Reproduction
• Areal Distribution
• Abundance
• Movements
• Predatory Biology
• Social & Complex
Behaviour
• Directed fishery threats
• Indirect mortality / beach meshing
• Habitat loss & Degradation
• Other problematic factors
• Past and present conservation
measures
• Action required & rationale
for CITES proposal
• Jaws
Carcharodon carcharias (Linnaeus, 1758)
Subclass
Elasmobranchii
Superorder Selachimorpha [Pleurotremata]
Order
Lamiformes
Family Lamnidae
Spanish: Tiburón blanco; jaquetón blanco; Tauró blanc
(Catalunya); salroig (Majorca); Marraco (Barcelona); Sarda
(Canaries)
Italian: Squalo bianco; pescecane; mangia alice;
damiano (Naples); tunnu palamitu di funnu (Catania); pici bistinu
(Messina)
Croatia: Pas ljudozder
Portuguese:
Tubarão
German: Menschenhai
Maltese: Kelb
il-bahar; Kelb-il-bahar abjad; Silfjun; Huta
tax-xmara
Afrikaans: Witdoodshaai
The
great white shark, within the monotypic genus Carcharodon, is
related to four other Mackerel sharks (Makos, Isurus spp. &
Porbeagles or Salmon sharks, Lamna spp.) in the Family
Lamnidae.
Arguably
equalled only by the killer whale (Orcinus orca) as a marine
macropredator, the great white shark occupies a cosmopolitan range
throughout temperate seas and oceans and will occasionally penetrate
tropical zones. Principally an epipelagic dweller of neritic waters,
Carcharodon is found from the surfline to well offshore, at the
surface and to depths over 250m on the bottom; commonly patrolling small
coastal archipelagos inhabited by pinnipeds , rocky headlands where
deepwater lies close to shore and offshore fish reefs, banks and
shoals.
Adult female specimens of Carcharodon
carcharias (Linnaeus, 1758), above; and below, the shortfin mako
Isurus oxyrinchus (Rafinesque, 1810). Note morphological
differences in respective medial fin positions and pigmentation in living
specimens. Juveniles of Carcharodon are regularly confused for the
shortfin mako. (Illustrations © Ian K. Fergusson).
White sharks vary from almost black to slaty-grey or dun above,
with the ventral surfaces predominantly white. A strong, variable line
of demarcation separates the dorsal and ventral surfaces with associated
blotching, particularly near the gill-slits and above the pelvic fin bases.
Small, irregular dark spots may be present on the flanks posterior to
the 5th gill slit and the lateral surfaces have a bronzy sheen that may
be structural and manifested only in strong sunlight. The ventral tips
of the pectorals are black and most specimens exhibit a black oval blotch
in the axil of the pectoral fin. The pelvic fins may be blotched with
olive or grey; likewise the ventral caudal lobe, with white marks and
white anterior booting. The mode of these demarcations vary greatly individually,
but may exhibit common trends amongst sharks in a given geographical remit.
The variance in dorsal pigment and tone is also very variable, especially
between centers of abundance but also within them. Californian examples
are often very dark slate-grey above, almost black. South African examples
(Cape Province) tend to be somewhat more dun or olive, as do Australian
specimens (but with much variance). Mediterranean specimens are predominantly
slate-grey or dark olive-brown dorsally.
The maximum size attained by white sharks remains a matter embroiled by
debate and spurious information. Adult females can reputedly attain
7 metres based on an unconfirmed specimen from Kangaroo Island, Australia,
taken in 1987. A further female taken off Filfla, Malta, in the same year
was reported as 713cm TL by local sources but can be demonstrated, on
the basis of a previously unexamined photo scrutinised in October 1998,
to be considerably smaller at circa 530cm TL. It is realistic to suggest
that maximum length of this species actually lies in the range of 550
to 600 cm TL. The smallest free-swimming examples, on the other hand,
appear to be ca. 120cm TL.
A recent allometric (size-on-size) equation for
weight (W) versus total length (TL) is given by Henry Mollet and Gregor
Cailliet (1996):
Until very recently, biological knowledge
of white shark reproduction was simply informed conjecture, based largely
upon comparisons with other better-studied Lamniform sharks such as sandtigers
and porbeagles. Records of pregnant white sharks in the literature were
effectively confined to a single Mediterranean report by Norman and Fraser
(1937), who discussed the capture of a 4.3 metre pregnant shark, reputedly
a great white, taken in the summer of 1934 off Agamy Beach at Alexandria,
Egypt. The shark contained 9 embryos, each about 61cm in length. Their
account raises some doubts over the accuracy of their original information
and perhaps the identification of the species. There was dearth of new
information until 1985, since when six pregnant females have been verified
and seven more reported. Pacific records make up the majority, and include
four from Japan: a 555cm great white from Kin, Okinawa, taken on February
16 1985; a ca. 470cm example from Taiji, Wakayama, on April 2 1986; a
480cm specimen off Uchinoura on May 14 1992 and a 515cm shark from Toyo-cho,
Kochi, captured a few days later on May 22 (Uchida et al., 1996). A 530cm
white shark containing 7 full-term foetuses was taken on November 13,
1991 at North Cape, New Zealand (Francis, 1996). As if to offer some contemporary
credibility to Norman and Fraser's report, the Mediterranean offered-up
a pregnant female in the shape of a five-metre-plus specimen taken in
a tuna net off Sidi Daoud, on Tunisia's Cape Bon, in the summer of 1992.
She contained two foetuses when eviscerated at the quayside, but may have
aborted more of a larger litter during capture (Fergusson, 1996). Further
recent but unconfirmed reports of pregnant white sharks originated between
1981 and 1996 from Queensland and South Australia. Three embryos of ca.100cm
were taken from a large great white caught near Taiwan in February or
March 1988, and off South Australia two embryos (one of 127cm) were found
during March 1994 in a net - probably aborted by the mother, which had
escaped, leaving a large hole. A very large female, reputedly of 640cm
TL and containing five embryos, was taken 5 miles off Malindi, Kenya,
in early August 1996 (Rodney Salm, IUCN, pers.comm) and is confirmed by
good photographs, although no photos of the litter have come to light.
Literature: Pratt and Casey (1985); Bigelow and Schroeder
(1948); Schroeder (1938, 1939); Springer (1939); Piers (1934); Scattergood
(1962); Scud (1962); Day and Fisher (1954); Templeman (1963); Guitart and
Milera (1974).
Literature:
Siccardi et al. (1981); Gadig and Rosa (1996); Gadig, pers. comm.
Literature: Quéro et al. (1978); Cadenat and Blache
(1981); Fergusson (1996); Ellis and McCosker (1991, for Azores).
Literature: Doderlein (1881); Lozano Rey (1928); Tortonese (1956);
Postel (1958); Bini (1960); Capapé et al. (1976); Fergusson (1995, 1996);
Mojetta et al. (in press).
Literature: Bass et al.
(1975); Cliff et al. (1989); Compagno (1991 and pers. comm); Compagno and
Fergusson (1994); Levine (1996); Smith (1951).
Literature: Cliff et al. (1989); Khalaf (1987);
L.J.V. Compagno (pers. comm.); R.V. Salm (pers. comm.).
Literature: Nakano and Nakaya (1987); Uchida et al.
(1996); Bruce (1992); Last and Stevens (1994); Francis (1996); Ayling and
Cox (1982); Cappo (1988).
Literature: Taylor (1985; 1993);
G. Burgess (pers. comm.).
Literature: Klimley
(1985); Royce (1963); Pike (1962); Fitch (1949); Kenyon (1958); Follet
(1966); LeMier (1951); Kato (1965); Bonham (1942).
Literature: Engaña and McCosker (1987)
Overall population estimates for this species are unknown and even regional
or localised estimates are sketchy. Inter-annual abundance of these sharks
can be very variable and unpredictable, giving rise to 'good' and 'bad'
years, in a colloquial sense, for great white numbers, albeit that the
causal factors are essentially unknown. After field observation and tagging
work, Rocky Strong et al. (1996) calculate the number of white sharks
frequenting the Dangerous Reef area of South Australia at circa 200 animals.
On a wider level, some landings data is available to (very roughly) indicate
the proportional abundance of white sharks, when compared to other species.
Game fishery captures of sharks in S.E. Australia between 1961 and 1990
indicated a catch-ratio of great whites of 1:22 in the 1960's, declining
to 1:38 in the 1970's and 1:651 in the 1980's (Pepperell, 1992). South
Australian game fishery catches from 1980-1990 averaged 1.4 sharks per
year and has declined since the 1950's, possibly through a reduction in
effort (Bruce, 1992). Off the eastern USA, National Marine Fisheries Service
statistics from 1965 to 1983 show a decline in ratio from 1:67 to 1:210
(Casey and Pratt, 1985). Other indices of catch-rates are available from:
a) California, between 1960-85 as 0 to 14 sharks per year (mean 3.2; Klimley,
1985); b) Natal, between 1974-88 as 22-61 sharks per year (Cliff et al.,
1989); c) Central Mediterranean Sea (Sicilian Channel), between 1950-94
as 0-8 sharks per year (mean 2.2, author, unpublished data). In other
areas such as Brazil and Hawaii, captures are much more nominal and sporadic
over time.
Occurrences are known within a wide sea-surface
temperature (SST) range of ca. 7.0 to 26ûC, but the species seems most
frequently encountered in temperate waters between 13 to 20ûC. Patterns in
movement and abundance within some areas (e.g., the seasonal position of
the 15ûC isortherm in the Northwest Atlantic [Casey and Pratt, 1985])
appear linked with seasonal variations in sea surface temperature, as
warmer waters influx poleward towards the northern limit of regional white
shark distribution. However, temperature may have only a limited effect on
the distribution of these sharks. An anomaly in white shark distribution,
at least on face-value, is their absence (or extreme scarcity) within
temperate coastal waters of Atlantic Europe - despite suitable habitats
and prey (including seals) being available to them.
Equally, it is clear that voyages by
white sharks can cover great distances. Although primarily occupying and
journeying within continental and insular shelf habitats, some white
sharks - and generally individuals from the larger size classes -
undertake long-distance journeys across the great ocean basins. They have
been found, albeit sporadically, both in mid-Atlantic at the Azores
archipelago and mid-Pacific at the Hawaiian Islands. Nominal captures have
also been reported from other discrete oceanic locales - including Gough
Island in the South Atlantic and Bikini Atoll in the Pacific's Marshall
Islands, as well as rather more captures from pelagic gillnetting
operations for squid in the epipelagic zone of the North Pacific. There is
no evidence to suggest that these treks have some reproductive
significance and the occurrence of white sharks at any one of these
localities is very variable in terms of both numbers, seasonality and bias
of gender. Taylor (1985, 1994) suggested that white sharks may frequent
Hawaiian waters through some predatory interrelationship with humpback
whale calving cycles. A similar hypothesis of a relationship between white
sharks and sperm whales may be applied to explicate occurrences at the
Azores, although in both cases this is very tentative. It is equally
likely that the phenomenon simply manifests the opportunistic, nomadic
behaviour of some larger white sharks - a trait that may conceivably
increase with older individuals that can readily subsist on large oceanic
bony fishes, cetaceans and other sharks. Whatever the case, these records
demonstrate that despite the wide geographic separation between known
centers of abundance, the white shark's distribution cannot be considered
as disjunct. For example, inter-hemispherical movement between temperate
areas, whereby white sharks traverse very warm equatorial zones, is
plausible. A possible mechanism is tropical submergence, where the shark
descends into and travels within deeper, cool oceanic waters across the
equatorial zone. However, white sharks also occur in shallow inshore and
offshore waters in the tropics and may may not need to use this mechanism
unlike some epipelagic and deep-water sharks.
So there appear to be
two sides of the same coin: the nomadic, 'just passing through' style of
spatial behaviour and the more systematic pattern of repetitive
occurrence, as demonstrated at select sites such as the Spencer Gulf, the
Farallons, and the Dyer Island Channel by tagging and field observation.
Clearly, the populaces of these 'classic' white shark localities are in a
state of continual flux, with numbers of sharks on any one day comprising
both 'resident' individuals and short-term nomadic visitors, and with all
of them moving around and seldom staying close to observers. It is
suspected that some white sharks occupy and patrol a favoured home range
in the broad sense of the word (particularly some large individuals at
Farallons), and that a proportion of attacks on divers are actually an
aggressive if limited response to the invasion of this 'personal space'.
The issue of white shark spatial behaviour is an enigmatic topic, whose
secrets may ultimately be revealed through current advances in electronic
tagging, data-logging tags, seabed-monitor arrays and long-distance
satellite telemetry..
The white shark is a formidible macropredator, with its primary
prey marine vertebrates and its most important prey bony fishes, cartilaginous
fishes, and marine mammals. Smaller sharks are primarily piscivorous,
but with growth the maximum prey size becomes larger and larger juveniles
and adults feed on a wide variety of small
to large bony fishes, elasmobranchs, cetaceans, and pinnipeds as their
principal prey, with other marine vertebrates and invertebrates also taken.
Fish prey recorded from across the size-range of this species includes
a number of elasmobranchs, such as juvenile dusky sharks, sandbar sharks,
tope, spiny dogfish, smoothhounds and other houndsharks, bronze whalers,
milk sharks, and blue sharks, shortfin makos, sandtigers, scalloped hammerheads,
whiptailed stingrays (Dasyatidae), batrays, eagle rays, guitarfish (Rhinobatidae),
wedgefishes (Rhynchobatidae), and elephantfish (Callorhynchidae). Bony
fish taken include bluefin tuna, Atlantic bonito, albacore, bullet tuna
and other scombrids such as frigate mackerel; hake, bluefish, swordfish,
sardines and other herring-like fishes, sturgeons, sea catfish, cabezon,
lingcod, rockfish, sea breams, croakers, jacks (Carangidae), barracuda,
striped bass, Pacific salmon, halibut and flounders, and even ocean sunfish.
Marine reptiles are sporadically ingested, and within Mediterranean waters
white sharks have been caught with the remains of Loggerhead Caretta
caretta and green turtles Chelonia mydas in their stomachs
(Fergusson et al., in press). Cephalopods (such as squid and cuttlefish),
gastropods and crustaceans are less important prey but benthic crabs may
be ingested in suprising numbers by large sharks, which don't necessarily
spurn small prey.
The role of Carcharodon as a primary predator upon marine mammals,
particularly pinnipeds, is oft-cited but sometimes overemphasised versus
the fish components in its diet. It is sometimes stated that larger sharks
switch to pinnipeds as prey and are narrowly dependent on them, but this
is apparently erroneous. In areas where pinnipeds are abundant and sympatric
with white sharks, such as along the coasts of California, the Cape Province
of South Africa,islands off South Australia, and the Otago region of New
Zealand, these sharks will lurk off haul-outs and prey upon a variety
of species including northern elephant seals Mirounga angustirostris,
fur seals Arctocephalus spp., harbour seals Phoca vitulina,
grey seals Halichoerus grypus, and sea lions including California
sea lions Zalophus californianus and Australian sea lions Neophoca
cinerea. The dynamics of such predator-prey relationships have been
the focus for recent Californian studies, primarily at the Farallon Islands
(Ainley et al., 1981; 1985; Klimley et al., 1992 and 1996) and at Año
Nuevo Island, a low, sandy feature north of Santa Cruz (Le Boeuf et al.,
1982; S. van Sommerman, pers.comm.). Similar field studies have been pursued
off Cape Province, South Africa, by the Shark Research Center (SRC) of
the South African Museum.
Cetaceans - and especially dolphins - are a common prey-item in the Mediterranean
and elsewhere, including within regions
where pinnipeds also occur and are preyed upon (e.g., Spencer Gulf of
South Australia; Bruce, 1992;Western Cape, South Africa). Taken free-swimming
and opportunistically scavenged from the nets of fishermen and beach meshing
programmes, dolphins are known to fall prey to great whites off South
Australia (Bruce, 1992), Western Australia (Cockeron et al.) South Africa
(Cliff et al., 1989; Compagno, 1991) and in the Mediterranean (Postel,
1958; Fergusson, 1994). In the latter area, bottlenose dolphins Tursiops
truncatus are the most frequent prey species. Other odontocetes (toothed
whales) taken by these sharks include common dolphins Delphinus delphis
off Tunisia (Postel, 1958), harbour porpoises Phocoena phocoena
in the Canadian Atlantic (Arnold, 1972), pygmy sperm whale Kogia breviceps
and Dall's porpoise Phocoenoides dalli off California (Long, 1992;
1996); also dusky dolphins Lagenorhynchus obscurus and Indo-Pacific
humpback dolphins Sousa plumbea within other regions. The carcasses
of great whales are readily scavenged by white sharks in feeding aggregations
off Long Island (e.g., Carey et al., 1982; Pratt et al., 1982), California
(P. Pyle, pers. comm; R. Collier, pers. comm), South Africa and elsewhere.
This interrelationship between white sharks and cetaceans is by no means
a new one - scars attributable to these sharks have been found marking
the fossilised skeletons of bottlenose dolphins (see Cigala Fulgosi, 1990,
for a detailed example from Italy) and cetotheriid whales (Demere and
Cerutti, 1982).
Interestingly, the white shark will often bite, but rarely ingest, two
groups of prey. Firstly, seabirds - particularly jackass penguins Spheniscus
demersus on islands off the South African coast - are commonly found
to have suffered cursory injurious or mortal 'grab-release' , 'slash'
(apparently with the tips of the lower and upper teeth), and 'bash' (with
the upper teeth) bites from white sharks (Randall et al., 1988; M.A. Marks,
pers. comm.) but are rarely ingested. Other avians including brown pelicans,
cormorants, gannets and gulls suffer similar fate, albeit not always culminating
in some disabling injury. Ian Fergusson witnessed an entire sequence at
Dyer Island wherein a cormorant (Phalacrocorax sp.) was grab-released
by a white shark, commencing with the surface-resting bird being taken
suddenly by a leaping white shark and carried airborne in its mouth, before
being taken subsurface for a few seconds. Much to his incredulity, the
bird then re-surfaced and flew-off, showing no visible sign of mishap
after the impromptu 'slam-dunk' (Fergusson, 1995). Birds are also 'bounced'
(sharks come up underneath gulls and other seabirds sitting on the surface
and send them tumbling with an upward flip of their heads), and are either
grabbed or not. Sea otters (Enhydra lutris) are mortally injured
by white sharks off central California, especially near Monterey and Carmel
(Ames and Morejohn, 1980; Ames, 1996) but have yet to be found in white
shark stomachs. Exactly why birds and sea-otters are almost exclusively
rejected as prey is unclear, but perhaps this phenomena has some enigmatic
bearing upon the reasons why most human victims of white shark attack
are also released, usually after a very limited and non-energetic contact
of a nature that is at odds with the unquestionable power of these fish
and the massively traumatic and swiftly lethal nature of full-fledged
feeding attacks on similar-sized pinnipeds and other marine vertebrates.
However, even regular prey animals such as young of the year Cape fur
seals may be lightly grabbed by white sharks and released with little
injury (M.A. Marks, pers. comm.).
White sharks will scavenge from fishermens nets and longlines and will
take all manner of hooked fish, including conspecifics, taken by rod-and-line.
This propensity often results in their own accidental entrapment. In Mediterranean
waters and elsewhere, it is because of this opportunistic behaviour that
many white shark captures are made by means of the predators scavenging
large fish and other sharks taken by longlines, or entrapped in gillnets,
on setlines or other fishing modes. In addition, large specimens are sporadically
taken by swordfish harpoons within the Straits of Messina, Sicily. Firm
evidence of these sharks scavenging fishery discard is afforded by the
discovery of a 2 metre bottlenose dolphin carcass in the stomach of a
535cm white shark taken at Favignana, Sicily, in May 1987. The dolphin
remains, bitten cleanly in two at the abdomen, were relatively fresh but
clearly scavenged - a tightly-knotted nylon rope was bound around the
caudal peduncle, an unmistakable sign that the mammal had been entrapped
by driftnet, drowned and then retrieved by fishermen. At the time, fearful
of environmental lobbying over cetacean mortality in these fisheries,
local operators would habitually sink dolphin carcasses by knotting rope
around the tails and attaching a stone anchor (Giuseppe Notarbartolo di
Sciara, pers. comm.). In all likelihood, this dolphin was ingested post-mortem
by a white shark which severed the carcass's anchor-rope during feeding.
Contrary to the lonely JAWS spectre of an idiot eating-machine, the white
shark is actually a social animal, exhibiting a raft of complex behaviours.
Intraspecific ethology and sociobiology in this species are now receiving
dedicated research attention, and are considerably more complex than previously
recognised or embodied within the archetypal 'lone killer' image. Field-observations
have described pecking-orders at feeding aggregations at carcasses which
seem based upon a size-hierarchy (see Pratt et al., 1982), where larger
sharks dominate in eating; this has also been seen off South Africa with
floating baits, but is complicated by individual motivation. Some of the
great white's swimming-modes are interpreted as ensuring avoidance of
conspecifics and maintenance of a personal space, such as cautiously-timed
'turn-aways' between two animals converging on reciprocal approaching
courses. Similarly, the 'parallel-swim' mode is often seen, whereby two
sharks heading on the same vector retain an unfluctuating distance from
each other, again as what seems to be maintenance of personal space (L.J.V.
Compagno, M.A. Marks and I.K. Fergusson; personal obs. and in press).
What appears to be a non-injurious means of deflecting competition whilst
feeding at the surface has been observed off California and South Africa,
whereby white sharks strike or splash conspecifics with the caudal fin
in a spectacular exhibition dubbed 'tail-slapping' (Klimley et al., 1996).
White sharks will also shove or 'body-slam' other white sharks (and boats)
by lateral movements of their bodies, and also use their caudal fins to
strike boats and even observers aboard them (M.A. Marks, pers. comm.).
Many white sharks, both adult and immature, have slash and puncture wounds
of superficial nature to the head and dorsum, which are inflicted by the
teeth of rivals during brief bouts of intraspecific aggression and competition,
especially near food resources but very likely also through courtship
or other social interactions. In the past, these marks were almost exclusively
explained as being caused by the flipper nails or teeth of pinniped prey,
but closer inspection and further observation in the field largely excludes
this theory. Aggression between these sharks is very inhibited, considering
the potential for severe wounding or worse, and includes rather cursory
bashing, slashing and grab-release biting with both the upper and lower-jaw
teeth. Where elicited on inanimate items offered to white sharks, much
of this casual mouthing seems to be investigative rather than any attempt
to ingest, and the observer is left with an overwhelming image of the
shark using its fearsome jaws with very fine control for tactile sense
and manipulation as well as for actual feeding (authors, personal obs.).
By the same token, such rather low-intensity biting is typical in attacks
on humans, with puncture marks, as opposed to actual flesh removal, being
commonplace. Whilst white shark attacks are often explained through a
process of 'mistaken identity' with natural prey, underwater observations
suggest that the sharks readily distinguish between humans and prey-animals
such as pinnipeds. A more compelling hypothesis is that many such 'hit
and run' interactions are motivated by interspecific aggression; perceived
invasion of personal space, or other motives not directly allied to feeding
such as 'play'. As white sharks apparently interact socially by low-intensity
biting and grabbing, at least some oral contact with humans and other
animals that are not regular prey may have a social framework, with sharks
behaving with humans as they would with other sharks. Humans in such interactions
may not understand what the shark's body language (including displays
such as gaping and hunching) may signify (if they notice them at all)
and what responses are appropriate to the context, until the shark proceeds
further in the interaction and grabs the person. Work in progress off
South Africa suggests that white sharks as individuals and groups confronted
by free-swimming skin and SCUBA divers are usually not aggressive even
when baits and chum-trails are present but may investigate the divers
quite closely without showing any signs of perturbed, agonistic behavior
(M. A. Marks, pers. comm.).
White sharks attracted in baited situations exhibit curious behaviours,
such as inverted surface-swimming with the mouth agape and gape-displays
on the approach to shark cages, divers and so-on, at least some of it
being thwart-induced (see Strong, 1996) or displacement behaviour, in
a generic sense, as well as int. This activity that is not unlike that
observed with the shortfin mako Isurus oxyrinchus. It may appear
simply maniacal on films, but such ethology must have a more deep-rooted
purpose. In the gape-displays, the palatoquadrate (upper jaw) is exposed
for a protracted period during non-feeding, rather akin to a dog snarling
and bearing its canine teeth. Indeed, its purpose may be exactly that
of the dog - an agonistic threat display that warns-off competitors or
intruders of personal space with a display of natural weaponry that requires
little further explanation! Some white sharks couple this type of behaviour
to a unnaturally stiff swimming mode, with pectoral fins held markedly
downwards, and back slightly arched ('hunch' display) and accompanied
by periodic gaping that may be directed both intraspecifically (to other
white sharks) and interspecifically (for example, to people - but quite
possibly, or probably, also to seals, dolphins and other animals). The
conspicuous dark axillar blotch (highlighted by bare white axillar skin
around it), which is only normally exposed by the 'pecs-down' hunching
style of swimming, may play a visual, communicative part in this posturing.
Notably, it is a feature of pigmentation found on both white sharks and
the shortfin mako (but not the porbeagle or salmon shark) - suggesting
that its role, if any, is duplicated or at least overlapping in both species.
It's occasional absence as a feature on Mediterranean white sharks may
be evidence of a low population density, with very limited intraspecific
contact; i.e., the feature has been progressively lost in an adaptorial,
evolutionary sense. However, white sharks without axillary spots also
occur off South Africa and California where white shark numbers are greater
and social contact injuries are commonly seen on sharks. Whilst this explanation
remains totally speculative, the apparently low degree of contact between
Mediterranean great whites other than for reproduction is further indicated
by the dearth of scratches, stabs and other allied marks as seen on the
heads and anterior parts of these sharks in regions elsewhere (see above).
Conservation of the great white shark
In directed terms, white sharks are primarily
exploited for the curio trade, with jaws and individual teeth fetching
high prices on both local and international markets; a recent large set
of jaws from South Africa being valued at $50,000. Often associated with
these commercial ventures are sportfisheries, as formerly undetaken in
a high-profile manner off Southern Australia, South Africa, and to a lesser
degree in other regions such as the eastern USA. Whilst a number of white
sharks are taken annually by rod-and-line charter-boats, a further, more
unscrupulous method employed by some operators involves passive-fishing
by means of large baited meathooks secured to anchored oil-drum floats
by heavy-gauge chains. These 'chain-line' and associated 'big hook' fisheries
are generally unstructured, unregulated or downright illegal; their aim
being solely the procurement of white shark jaws or teeth for trade. Individual
dentition removed from smaller white sharks may sell for a higher price
in aggregate than complete sets of small jaws, and the threat posed by
the curio-trade encompasses white sharks across the entire size and maturity-range.
This trade is not restricted to jaws and teeth; even entire specimens,
some attaining in excess of 5 metres, have been preserved by freezing
or taxidermy for static public display or as private trophies and curios.
In some areas, the excellent, palatable flesh of this species, rich in
red muscle when properly filleted, can fetch high retail prices for human
consumption and considerably more than that of other similarly-exploited
sharks such as the shortfin mako Isurus oxyrinchus. However, coupled
equally to consumer disquiet over the 'maneater' tag, and the fact that
white shark meat is not uncommonly high in mercury, commercial use for
human cuisine is geographically and culturally patchy. Because very few
specimens marketed by fisheries are listed under species-specific landings
statistics, consequently the precise level of such exploitation remains
poorly documented.
Some rod-and-line sports fisheries have operated both on the Atlantic
and Pacific coasts of North America, but rarely are these directed solely
at white sharks. In the Mid-Atlantic Bight, juveniles are occasionally
landed in angling tournaments - some 140 specimens were taken in those
waters between 1971 and 1991, by tournaments and research vessels (Pratt,
1996). In the past, big-game charters were run with some success to target
adult specimens in the same area. One operator, Frank Mundus of Montauk,
Long Island, was particularly active in this regard and his association
with catching big white sharks inspired the 'Quint' character in Peter
Benchley's JAWS. More opportunistic ventures, such as the harpooning of
sharks feeding on whale carcasses, was not unknown in the waters off Long
Island. The National Marine Fisheries Service (NMFS) now seeks to control
and outlaw targeted captures of white sharks in the northwest US Atlantic
(now supported by legislation enforced since early 1997) and has successfully
encouraged and coordinated the tagging and releasing of these and other
species of sharks for many years.
On the US Pacific coast, white sharks have been targeted in the past off
both central and southern California, albeit with variable success. Particularly
in the wake of "Jaws" (1975), white sharks were a popular but somewhat
elusive quarry for anglers in Southern Californian waters. Their activities
were frequently inspired by the large cash rewards offered at the time
by oceanariums, who sought great whites (or bits and pieces thereof) for
public display as a cash-in on the 'Jaws-mania' of the era. Juvenile sharks
were often taken very close inshore and larger specimens fell victim to
harpoon-wielding hunters. More recently, in the mid-1980's, one-off opportunistic
ventures were undertaken to capture white sharks at the Farallon Islands
near San Francisco, a National Marine Sanctuary reknowned internationally
as a white shark locality. The virtually simultaneous capture of four
sharks at the islands in 1984 was met with dismay by the scientific establishment
and some sectors of the public and media. Since then, fishing-forays to
the islands or elsewhere in Californian waters in search of white sharks
have been outlawed by State Legislation SB-144.
The propensity for white sharks to
scavenge from commercial longline and net fisheries makes them very
vulnerable to accidental entrapment. Worldwide, specimens are reported
annually from gill-nets, trammels, herring-weirs, purse-seines,
tuna-enclosures and hooked upon surface and bottom longlines and
set-lines.
Being a highly mobile, pelagic species the threat of habitat loss may
appear generally minimal to white sharks. However, disruption of prey
availability poses potential threats and in at least one area - the north
Adriatic Sea - white shark captures and sightings have declined markedly
in the latter half of this century, possibly through depletion of teleost
prey stocks, dolphins and possibly monk seals. All of these factors may
themselves be the effects of increased pollutants, overfishing or both.
Despite this, white sharks are adaptable predators capable of shifting
diet as conditions dictate and may simply cease inhabiting a degraded
area and move elsewhere. As tagging studies have demonstrated, these sharks
can display a varying degree of localised residency - a trait that may
make them vulnerable to habitat loss, at least within the short-term.
In recent years, cage diving for white sharks has become an important
'spot' industry in some parts of the world, most notably in Australia's
Spencer Gulf (Dangerous Reef and Neptune Islands), along the South African
Cape Coast (especially at Dyer Island) and to a lesser degree in the warm
waters of Baja California.
The most significant problem in applying definitive conservation
measures in favour of white sharks remains the lack of baseline data on
basic parameters such as the fecundity, age and growth and population-numbers
of the species. Such data is particularly difficult to obtain for white
sharks, principally as they are generally rare wherever they occur; esentially
cryptic, and pregnant females are scarcely captured let alone examined
by fisheries workers. Recruitment-rates for this species are unknown and
tagging studies, until recently involving unsuitable methodology for white
sharks, provides data that in this species takes a considerable number
of years to amass as a consequence of rather nominal recaptures and tag-returns.
Current topical thought, therefore, is that protective measures should
be based on a precautionary principle, until such time as more biological
information has been collated.
Protection in South Africa
It was acting precisely upon such guidance that the South African Government
legislated to protect white sharks in April, 1991 - the first nation to
do-so. The crucial points that convinced Louis Pienaar, then Minister
of Environmental Affairs, to create a ban on the sale of white sharks
or any white shark product, were a number of arguments proposed in a comprehensive
review by Dr Leonard J.V. Compagno (Compagno, 1991a) and others, strongly
supported by Nan Rice of the Dolphin Protection Group (who was instrumental
in lobbying Environmental Affairs to protect the white shark), which was
circulated prior to a meeting convened by Environmental and Sea Fisheries
agencies on March 19th of that year. On the basis of the report, the parties
had essentially agreed on total protection by the time of the meeting.
It was stressed that: "....Because the population size is likely to be
small, this shark could be threatened if exploitation by trophy-hunters
increased, as appeared to be the case, judging by the number of foreign
enquiries about commercial shark hunting safaris into Cape waters....The
reproductive rate is low and the slow growth and late maturity of this
species makes it highly vulnerable to over-exploitation..." (see Smale
[1991] for a full summary, and Compagno, 1990, 1991a,b, for particulars
of white shark protective measures in South Africa). The legislation concerning
protection of this species falls under the National Marine Fisheries Act
and covers the entire South African coastline and EEZ. It mandates heavy
fines, boat confiscation, and jail sentences for anyone caught catching,
killing or injuring white sharks, or dealing in white shark products.
Further legislation passed in 1998 tightens access to white sharks after
problems were identified, with research and ecotouristic shark cage diving
by permit only and with a limited entry for cage dive operators nationally
and for favored sites such as the Dyer Island Channel. Apparently the
South African protective action inspired the Namibian goverment to adopt
similar protective measures for the white shark in 1993.
Protection in California
Similarly, a set of protective measures were installed for Californian
waters, after lobbying by scientists and conservationists who found a
political ally in Sacramento Assemblyman, Mr Dan Hauser. Passed unanimously,
State Assembly Bill AB522 was enacted on January 1st, 1994 and effectively
prevented the development of directed sports or commercial fisheries for
white sharks in California until at least 1998. It is instructive to note
that the legislation received unequivocal support from, amongst others,
the California Urchin Divers Association and Surfrider Foundation, both
whose members figure in white shark attack statistics. Current legislation
concerning protection of this species is State legislature SB-144 (1997)
which replaces the temporary AB-522 enacted in 1994 and includes a clause
to outlaw all directed efforts at attracting white sharks either by use
of baits, chum, or other methods.
Protection in Eastern United States waters
After California's action, the State of Florida followed-suit with unilateral
state legislation, However, in early 1997, all directed targeting of white
sharks other than tag-and-release sport angling was outlawed throughout
all US eastern coastal waters and the Gulf of Mexico as part of the Fisheries
Management Plan (FMP) enacted by the government's National Marine Fisheries
Service (NMFS).
Protection in Australian Commonwealth waters
Despite the considerable efforts by many individuals there, Australia
had been slow to act in affording full protection for white sharks but
an increasing movement of public shark-sympathy, supported by a number
of scientific and conservation bodies, ensured that it was a matter never
far from media attention. Although commercial fishing still presents the
greatest threat to sharks off Australia, it had been sportfishing - and
especially for great whites - that commanded the greater media and public
attention. The South Australian towns of Port Lincoln and Whyalla, on
the shores of the Spencer Gulf, have long been traditional locations from
where big-game anglers set-sail in search of the ultimate fishing trophy.
And yet the sport was under increasing threat as the mid-1990's approached,
as evidence by frequent public wrath at reports of white shark captures
published within the Australian newsmedia.
In May 1995, Jon Presser and Ross Allan (1995) gave a discussion paper
suggesting a series of conservation measures for white sharks in South
Australian waters, including prohibition of chumming within 2 nautical
miles of the coast, restriction on fishing hook sizes, licensing of cage-viewing
tours and the expansion of conservation zones in the Spencer Gulf to restrict
possible access to white sharks. Central to these measures was the proposed
full protection of white sharks, under Section 42 of the Fisheries Act,
1982. Meanwhile, more 'unilateral' precautionary protection to white sharks
had been enacted by the States of New South Wales, Tasmania, Queensland,
Western Australia and - at least in a partial sense - by South Australia.
Through a joint press release issued on 22 December 1994, the Minister
for Primary Industries and Minister for Environmental Affairs acknowledged
the need to provide greater protection for white sharks and requested
recommendations from the research community. Final collation of this data
was completed on July 28, 1995, with a view to legislation being in-place
before the end of the millennium.
On 17 December 1997, Federal Environment Minister Robert Hill announced
that the Australian Government was protecting great white sharks throughout
all of their Commonwealth waters, prohibiting all directed hunting, molestation
or harassment of the animals. Anyone wishing to capture a specimen for
'scientific purposes' must hold a government permit. The white shark is
currently listed as a Federal Endangered Species. The State of Victoria
enacted local species-specific legislation in 1998; meanwhile, the Australian
government has agreed to pursue a CITES proposal for the white shark,
to which various parties and individuals are currently (Dec. 1998) offering
input.
Although we know very little about
the global interrelationships of these sharks, it may be the case that
regional or national protective measures will not ultimately address the
long-term welfare of the species. Citing the west coast of North America
by way of example, the white shark can be assured of total conservation
only if Oregon, Washington and Mexico followed California's lead. The
potential ineffectiveness of unilateral measures is perhaps best envisaged
when one considers a white shark pupping and nursery-ground - the Sicilian
Channel. If measures were enforced solely in Sicilian (Italian)
jurisdiction, little effect would be manifested in the protection of
sharks only a few kilometres distant, in the seas off Tunisia.
Ultimately, the white shark's future can only be secured
through legislation enacted and enforced pan-globally. Thus, seeking a
listing within the remits of CITES protection is considered imperative
considering the wide-ranging habits of this fish.
Directed fishery
exploitation (mortality) of white sharks is primarily undertaken with the
aim of securing and trading its body-parts, especially teeth and jaws, as
trophies or curios. Recent information suggests that preserved jaws may
fetch from $10,000 - $50,000 in collectors circles, and there is good
evidence from a Scottish-based collector_ that white shark teeth and jaws
are still freely available from California and South Africa despite the
current trade embargos there. Anecdotal evidence supports the notion of
unlawful trade being undertaken in the latter nation's Cape Province
waters, with local commercial fisheries operations supplying international
offshore fishing vessels with white shark jaws and shark cage operators
being offered enormous sums for white shark jaws and isolated teeth.
Similarly, new suppliers have offered collectors white shark jaws from an
offshore fishery based at Mazara del Vallo (Trapani), Sicily, which
incidentally captures neonatal specimens. Some South American states are
emerging as a secondary source of collectable material, apparently pooled
from sharks caught off Baja and environs, where Mexican-based dealers act
as couriers.
The vulnerability of white sharks to curio-hunting is
poignantly comparable to previous exploitation of large terrestrial
predators for trophies, furs and so-on. All available evidence points to
these key facets in proposing the white shark for CITES listing:-