Review of the great white shark
Carcharodon carcharias

Prepared by Ian K. Fergusson,
Shark Trust & IUCN Shark Specialist Group

(Conservation section partly based on the IUCN-SSG's Red List species account, prepared by I.K. Fergusson, L.J.V. Compagno and M.A. Marks)

Copyright © 1998. All rights reserved, No reproduction permitted in any form without the express permission on the author.

 

Contents

Status & Biology:
Nomenclature & Taxonomy
Description
Size
Weight-Length relationship
Reproduction
Areal Distribution
Abundance
Movements
Predatory Biology
Social & Complex Behaviour

Conservation:
Directed fishery threats
Indirect mortality / beach meshing
Habitat loss & Degradation
Other problematic factors
Past and present conservation measures
Action required & rationale for CITES proposal


Great white shark
Carcharodon carcharias (Linnaeus, 1758)

Class Chondrichthyes
Subclass Elasmobranchii
Superorder Selachimorpha [Pleurotremata]
Order Lamiformes
Family Lamnidae

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)
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

Current IUCN Status Assessment: VULNERABLE globally

Taxonomy & Early History
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.

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.

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• Status and Biological Summary

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.

Description

Carcharodon carcharias,  female (© Ian Fergusson)


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).

Isurus oxyrinchus, female (© Ian Fergusson)

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:-

13 / 11-12

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
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.

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Size and Reproduction
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.

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
A recent allometric (size-on-size) equation for weight (W) versus total length (TL) is given by Henry Mollet and Gregor Cailliet (1996):

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

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• Reproduction
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.White shark dorsal fin (© I. Fergusson)

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.

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Distribution

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).

"Great white shark under Mediterranean sun", by Ian Fergusson

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).
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).

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).
Literature: Siccardi et al. (1981); Gadig and Rosa (1996); Gadig, pers. comm.

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.
Literature: Quéro et al. (1978); Cadenat and Blache (1981); Fergusson (1996); Ellis and McCosker (1991, for Azores).

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).
Literature: Doderlein (1881); Lozano Rey (1928); Tortonese (1956); Postel (1958); Bini (1960); Capapé et al. (1976); Fergusson (1995, 1996); Mojetta et al. (in press). See Bibliography for detailed listing.

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.
Literature: Bass et al. (1975); Cliff et al. (1989); Compagno (1991 and pers. comm); Compagno and Fergusson (1994); Levine (1996); Smith (1951).

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).
Literature: Cliff et al. (1989); Khalaf (1987); L.J.V. Compagno (pers. comm.); R.V. Salm (pers. comm.).

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).
Literature: Nakano and Nakaya (1987); Uchida et al. (1996); Bruce (1992); Last and Stevens (1994); Francis (1996); Ayling and Cox (1982); Cappo (1988).

Central Pacific:- Hawaiian Islands off Oahu, Hawaii and Northwestern Islands (R); Marshall Islands at Bikini (VR); Easter Island, based on recent shark attack.
Literature: Taylor (1985; 1993); G. Burgess (pers. comm.).

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).
Literature: Klimley (1985); Royce (1963); Pike (1962); Fitch (1949); Kenyon (1958); Follet (1966); LeMier (1951); Kato (1965); Bonham (1942).

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.
Literature: Engaña and McCosker (1987)

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• Abundance
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.

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• Movements and Nomadicy
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.

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.).White shark towing 'Crittercam' equipment (© I. Fergusson)

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.

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..

 

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Predatory Biology
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 feedWhite shark breaching in False Bay; © African Shark Ecotours 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,White shark attacking Cape fur seal; © I. Fergusson 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.

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Social and Complex behaviour
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).

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Conservation of the great white shark

Threats

• Directed Fisheries
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.

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• Indirect Mortality and Beach Meshing
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.

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.Juvenile white shark, Tunisia, 1998

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.

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• Habitat Loss or Degradation
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.

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•Other Factors
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.

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 (see News piece for fuller description).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.

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Conservation measures

• Present and Planned action
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.

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Action Required

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.

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.

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. White shark, Sicily

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?
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:-

  • The species is specifically targeted for trophies, saleable curios, and fins for the oriental soup-fin trade. Current unilateral restrictions of such activities, as exist in some parts of the species' range, is presently the only curb upon such exploitation. The effectiveness of such legislation is wholly reliant upon diligent enforcement in those nations or states where it has been enacted. Consequently their remains the potential for illicit trade and unlawful fisheries to continue operating, not least fuelled by the high prices paid by collectors for white shark jaws (as much as $10,000 dollars per pair of jaws and with collectors apparently willing to pay far more for jaws from large adults) and isolated teeth. Fin sets from large white sharks are valuable for their size, being priced at hundreds or even thousands of dollars, but also from their source, as white shark notoriety apparently boosts values of their fins.
  • There is growing evidence to suspect a global decline of white shark populations due to a combination of causal factors which are poorly understood, but which may include wide-ranging and largely unreported bycatch from world fisheries (including major offshore operations), as well as targeted fisheries. As the interrelationships, interchange and recruitment between seemingly discrete populaces of this shark are unknown, it is quite possible that regional mortalities have a much wider knock-on effect in a global sense.
  • Some areas where white sharks are periodically commercially targeted for trophies, or have been so in the recent past, also serve as preferred feeding habitats, nursery zones or other areas (white shark 'sites') and are believed important to reproductive behaviour and to social interactions. The sharks readily investigate boats at these white shark sites, and can be lured to within touching distance by chum and floating baits. As a result, white sharks are rather easily located and can be killed with rather modest effort and equipment (including relatively small outboard 'skiboats') in comparison to their value.
  • Given the wide areal range of white sharks, the limited geographical protection as currently exists in South Africa, California, eastern USA, Australia and elsewhere will not prevent dedicated exploitation shifting to other vulnerable locations where such hunting is not outlawed and white sharks are accessible.
  • Geographically-limited or unilateral restrictions will not ultimately curb the worldwide collectors market for white shark products. There is evidence that existing laws are already being violated, as discussed above.

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...

© Ian K. Fergusson 1998
No Reproduction in any form without express permission of the author.

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