Iles Crozet (Crozet Islands)
The Iles Crozet (Crozet Islands) are an archipelago in the south Indian Ocean, about two-thirds of the way from Madagascar to Antarctica, and part of the French Southern and Antarctic Lands. They are known for their wildlife, including fur seals, leopard seals, and southern elephant seals, as well as numerous bird species.
A large archipelago formed from the Crozet Plateau, Iles Crozet is divided into two main groups, bisected by the 2km deep Indivat basin:
L'Occidental (the West), which includes:
Île aux Cochons is the third largest island, at 70km2. This island is a 600m high volcanic cone, and its open crater consists of alternations of pyroclastic rocks and lava flows.
Ilots des Apotres is 2km2 in area, and reaches heights of 289m,
Ile des Pingouins is 3km2 in area, and reaches heights of 420m
the reefs Brisants de l'Heroine
L'Oriental (the east), which includes:
Île de l’Est is the second largest island in this archipelago, with an area of 120km2 . The sheer cliffs on this island reach 1100m high. This deeply dissected shield volcano is made up of two basalt units, one basal and one upper, the two separated by a pyroclastic horizon.
Île de la Possession is the largest island, with an area of 130km2. It reaches a height of 934m.
Discovered and claimed by France in 1772, the islands were used for seal hunting and as a base for whaling.
The two eastern islands are the oldest among the Crozet islands, at approximately 8 million years old. Of the western islands, Île aux Cochons is 0.4 million years old, Îles des Pingouins is 1.1 million years old and Îles des Apotres is 5.5 million years old.
The archipelago experiences a cold, oceanic climate, with an annual mean temperature of 5ºC, monthly mean temperature ranges between 2.9ºC and 7.9ºC in winter and summer respectively, an annual rainfall of 2400mm per year, and strong winds.
Originally administered as a dependency of Madagascar, they became part of the TAAF in 1955.
The islands are uninhabited except for 18 to 30 people staffing the Alfred Faure research station on Ile del la Possession.
Geographic Coodinates: 46 25 S, 51 00 E
Area: 352 sq km
Maritime claims: territorial sea: 12 nm
Climate: windy, cold, wet, and cloudy
Terrain: a large archipelago formed from the Crozet Plateau is divided into two groups of islands.The highest point is Pic Marion-Dufresne (1,090 m).
Natural Resources: fish, crayfish
Land Use: 100% tossock grass, heath, and fern
- introduction of foreign species on Iles Crozet has caused severe damage to the original ecosystem
- overfishing of Patagonian toothfish around Iles Crozet
The Iles Crozet (Crozet Islands) are included within the Southern Indian Ocean Islands tundra ecoregion defined by the World Wildlife Fund which encompasses five island groups in the southern Indian Ocean: Prince Edward, Crozet, Kerguelen, Heard, and McDonald Islands.
Endemic vascular plants include Poa cookii and Pringlea ascorbutica (both found on all the islands in the ecoregion). P. ascorbutica is the famous Kerguelen cabbage that sailors used to eat to prevent scurvy in days past.
Other endemic vascular plants include Polystichum marionense (only found on the Crozet and Prince Edward Islands), Ranunculus moseleyi (only found on the Prince Edward, Crozet and Kerguelen Islands), Poa kerguelensis (only found on Heard and the Crozet Islands) and Colobanthus kerguelensis (possibly on all the islands).
Other non-endemic, but notable, plant species include Crassula moshata, Aceana magellanica, the cushion-shaped Azorella selago, the feathery Cotula plumosa, and the grass Agrostis magellanica. Among the non-flowering plants, the fern Blechnum penna-marina should be noted.
A number of lichen species are endemic to the ecoregion, but many lichens in this ecoregion are bipolar. Many liverworts are endemic to the ecoregion. Plant distribution patterns indicate a strong relationship between the ecoregion islands, a relationship that has been found in some invertebrate groups as well.
The first thing a casual observer notices about this ecoregion is the overlap of normally sub-Antarctic and normally Antarctic species, from plankton, to fish, to higher predators. One therefore finds on these islands species that may be best suited to one or other of the two environmental regimes, but not necessarily their combination. For instance, the breeding success of the gentoo penguin (Pygoscelis papua) is lowest at its northernmost breeding localities (the Prince Edward and Crozet Islands). This may be because at these relatively northern latitudes the species has to hunt less suitable sub-Antarctic marine prey.
Three of the most obvious sympatric congeneric species in the ecoregion are the two sooty albatrosses (Phoebetria spp.), the two giant petrels (Macronectes spp.) and the two fur seals (Arctocephalus spp.). The sooty albatross (Phoebetria fusca) and the light-mantled sooty albatross (P. palpebrata) breed sympatrically only within this ecoregion i.e., on Marion, the Crozet and the Kerguelen Islands. P. palpebrata, the longest-lived of all birds, normally has a more southerly, Antarctic range than does P. fusca. P. palpebrata from Marion Island, for instance, must therefore fly further south than its congeneric to forage. This places more stress on this species at that locale. In contrast to Marion Island, where its population is much larger than that of P. palpebrata, P. fusca is outnumbered by its congeneric on the Kerguelen Islands. The Kerguelen Islands are, in turn, the most southerly breeding point of the more northern species. Interspecific competition for food and breeding sites between these two congenerics is low or non-existent where they breed sympatrically, and there are no recorded attempts at interbreeding.
However, at times sympatric congenerics do attempt to interbreed. The Northern Giant petrel Macronectes halli and Southern Giant petrel M. giganteus breed sympatrically within the ecoregion, although their sympatric breeding area spreads over a wider range outside of it. Burger and Johnstone reported failed breeding attempts between these two species on Marion and Macquarie Islands respectively (the latter island outside the ecoregion). Although Johnstone et al. remarked that, on islands where sympatric breeding occurs, there is genetic isolation between the two species, they point out that on Gough Island (also outside the ecoregion), Macronectus spp. individuals show a combination of both species’ characteristics. This suggests the ability to interbreed at some sites, although the reasons behind this may be only locally relevant.
The only two eared seals (otarids) in the region, the antarctic fur seal (Arctocephalus gazella) and the subantarctic fur seal (A. tropicalis), breed sympatrically only on the Prince Edward, Crozet and Heard Islands within the ecoregion, and on Macquarie Island outside of it. Only a small percentage of the worldwide population of A. gazella is found in this ecoregion, the Antarctic fur seal being otherwise exclusively confined to south of the Antarctic Convergence. Outside the ecoregion, A. tropicalis breeds at Amsterdam, Saint Paul, Gough and Tristan da Cunha Islands, all to the north. Interbreeding between A. gazella, which normally breeds south of the convergence, and A. tropicalis, which normally breeds north of it, has been reported from islands where they breed sympatrically. The possibility that the Antarctic fur seal may be limited by its Antarctic adaptations on the more temperate climate of Marion Island has been mooted. Bester and van Jaarsveld have also reported that the sub-Antarctic fur seal is smaller on Marion Island than it is at more northern latitudes. There are no indigenous land mammals throughout the terrestrial sub-Antarctic.
A number of birds that breed in the ecoregion have been classified as "Vulnerable". These are:
- Eaton’s pintail (Anas eatoni) (an endemic),
- wandering albatross (Diomedea exulans),
- sooty albatross (Phoebetria fusca),
- Indian yellow-nosed albatross (Thalassarche carteri),
- gray-headed albatross (Thalassarche chrysostoma),
- rockhopper penguin (Eudyptes chrysocome),
- macaroni penguin (Eudyptes chrysolophus),
- southern giant petrel (Macronectus giganteus) and
- white-chinned petrel (Procellaria aequinoctialis).
Macronectus giganteus and Eudyptes chrysocome may possibly qualify for "Endangered" status, while Phoebetria fusca may even qualify as "Critically Endangered".
Lower risk, but "near threatened" bird species found in the ecoregion are Kerguelen tern (Sterna virgata) (an endemic), northern giant petrel (Macronectus halli), the gray petrel (Procellaria cinerea) (one of the winter breeding petrels), gentoo penguin (Pygoscelis papua), light-mantled albatross (Phoebetria palpebrata) and black-browed albatross (Thassalarche melanophrys). All albatrosses breeding in this ecoregion have been accorded "vulnerable" or "near threatened" status. Albatrosses have "the highest proportion of threatened species in any bird family that comprises more than a single species".
Other than those species already mentioned above, the following birds also breed in the ecoregion:
- sub-Antarctic skua (Catharacta skua);
- kelp gull (Larus dominicanus);
- fulmar prion (Pachyptila crassirostris),
- Antarctic prion (Pachyptila desolata),
- thin-billed prion (Pachyptila belcheri),
- fairy prion (Pachyptila turtur),
- Antarctic tern (Sterna vittata);
- cape petrel (Daption capense),
- black-bellied storm petrel (Fregetta tropica),
- gray-backed storm petrel (Garrodia nereis),
- blue petrel (Halobaena caerulea),
- Wilson’s storm petrel (Oceanites oceanicus),
- common diving petrel (Pelecanoides urinatrix),
- South Georgian diving petrel (Pelecanoides georgicus),
- white-headed petrel (Pterodroma lessonii) and
- soft-plumaged petrel (Pterodroma mollis).
Only one group of oceanic islands, the Crozet group, supports a greater number of breeding seabirds than the Prince Edward Islands. All six species of albatrosses that breed in the ecoregion breed sympatrically on the Crozet Islands. 80% of the world’s population of Salvin’s prion (Pachyptila salvini salvini) is found on Île de l’Est of the Crozet group. Certain wandering albatross (Diomedea exulans) populations outside the ecoregion may not be of the same subspecies as those within the ecoregion, which, if confirmed, will mean that the breeding range of by far the majority of the ecoregion’s subspecies is confined to that area. More than half the world’s population of king penguin is found on the Crozet archipelago. The world’s largest colony of this highly specialized mesopelagic predator is found on Île aux Cochons of the Crozet group, with 300,000 breeding pairs.
The king penguin (Aptenodytes patagonicus) is unique among penguins, in that it has a breeding cycle that takes longer than a year to complete. The two major predators of the king penguin are the sub-Antarctic skua (Catharacta skua), which takes eggs and small chicks, and the southern giant petrel (Macronectes giganteus), which can take up to 10% of the chicks. The two sibling petrel species, the northern giant petrel (Macronectes halli) and southern giant petrel (M. giganteus) are the largest predator/scavengers in the Southern Ocean ecosystem. Apart from their normal prey of penguins, burrowing petrels, seal carrion, cephalopods, crustaceans and fish, Punta and Herrera have observed them preying on adult imperial cormorants in Argentina, and they have been observed to seasonally access crustacean prey items from the stomachs of their fish and penguins kills at Kerguelen and Crozet Islands, leaving the rest of the original prey item untouched. There is even a recorded instance of a M. giganteus individual killing and partly eating an immature black-browed albatross (Thalassarche melanophrys) off Newfoundland Head (outside the ecoregion).
There may be competition for food between the subantarctic fur seal (Arctocephalus tropicalis) and the southern elephant seal (Mirounga leonina) at certain sites within the ecoregion, for example Marion Island. Both fur seals are increasing in numbers throughout their ranges since sealing ceased, and at Crozet it is reported that both species’ populations are reaching their maximum possible growth rate. These seals were once driven to the brink of extinction by hunting. Interestingly, there exists a large breeding colony of the Antarctic fur seal (A. gazella) on Îles Leygues of the Kerguelen group that was never harvested, which recolonized the main island after exploitation had ended.
Southern elephant seal individuals also move between the Crozet and Prince Edward Islands, but no movements between the Crozet and Kerguelen Islands have been recorded, suggesting that the two populations are isolated from one other. Previous steep population declines of M. leonina at Marion, Prince Edward, Crozet, Kerguelen and Heard Islands appear to have now ended.
Among other suggestions, the previous population decline of Mirounga leonina at the Crozet Islands was linked to killer whale (Orca - Orcinus orca predation). A particular pod of O. orca at the Crozet Islands has developed an interesting hunting technique whereby individuals intentionally strand themselves on the shore in order to capture prey. Juveniles are taught to strand themselves by specific, often unrelated members of the pod that specialize in such teaching duties. Once the juveniles have mastered this skill, they must still be helped back to the sea with their prey by older members for their first few hunts. Of the other cetaceans associated with this ecoregion, the sei whale and fin whale populations around the Prince Edward, Crozet and Kerguelen Islands have been judged to be from two single stocks.
Types and Severity of Threats
Plant and/or animal aliens have invaded all the islands in the ecoregion, apart from McDonald Island (as far as is known). Some of these invasions have had serious effects.
Some of the most prominent alien plants on one or more of the islands of the ecoregion are Agrostis stolonifera, Poa annua, Poa pratensis, Cerastium fontanum, Rumex acetosella, Sagina apetala and Stellaria medea. Alien plants can, and do, have different effects. Poa annua is an alien plant that has colonized most sub-Antarctic islands.
Attempts have been made to introduce salmonid fish into the rivers of Marion, Crozet, and Kerguelen Islands. The introductions have only been successful on Kerguelen, which has relatively large rivers.
On the French Islands, many mammalian invaders are present, five of which were deliberately introduced (cats, rabbits, reindeer, sheep and the Corsican mouflon).
The European rabbit (Oryctolagus cuniculus) occurs on both French archipelagos, to which it was introduced in the 1800s. It destroys indigenous vegetation (e.g. Pringlea antiscorbutica and Azorella selago), as well as the reproduction sites of burrowing petrels. The main effect of the O. cuniculus is indirect, though, as it provides a food source for cats during winter. This allows the cat population to remain relatively high outside the bird breeding season, resulting in a higher impact on the avifauna the next year.
The two rodents, the house mouse (Mus musculus) and the black rat (Rattus rattus) were both introduced to the Crozet archipelago in the 1800s. Mus musculus was introduced to the Kerguelen archipelago at around the same time as it was at Crozet, but Rattus rattus only found its way to the Kerguelen archipelago in the 1900s. On Crozet, Rattus rattus preys on the chicks and eggs of small petrels, and may have been responsible for the disappearance of the blue petrel (Halobaena caerulea), the gray-backed storm petrel (Garrodia nereis) and the Kerguelen diving petrel (Lugensa brevirostris) on Île de la Possession, to which island the rat is confined. Other petrel species have shown reduced survival on this island. However, those species that are able to breed above 400 meters (m) at Île de la Possession, can do so in large numbers. On Kerguelen, it has been noted that rats are mostly concentrated near the human settlement, and therefore cause little damage to the avifauna there.
Île aux Cochons is the only island of the Crozet archipelago to which Felis catus has been introduced. Felis catus was introduced to this island before 1887, and has subsequently caused the local extinction of the white-chinned petrel (Procellaria aequinoctialis), the gray petrel (Procellaria cinerea), and the gadfly petrels. Felis catus has been specifically linked to the "vulnerable" status accorded the endemic Eaton’s pintail (Anas eatoni), which occurs only on these two French archipelagos (see Table 1). It does not enter the burrows of small petrels, as they are too narrow, and must instead rely on capturing this prey when it arrives or leaves the colony. This allows the smaller Salvin’s prion (Pachyptila salvini), fairy prion (Pachyptila turtur) and South Georgian diving petrel (Pelecanoides georgicus) to remain numerous on Île aux Cochons.
Chapuis et al. best describe the situation on the French Islands. "The three main rules for introductions defined by Pimm have been broken:
(1) introduction in the absence of predators and competitors;
(2) introduction of highly polyphagous species (herbivores);
(3) introduction in ‘relatively simple communities where the removal of a few plant species will cause the collapse of entire food chains.’"
However, this commentary on the modification of the French territories must conclude by noting that some of the smaller islands within these two archipelagos remain unmodified by alien mammals. The environments of many of these smaller islands have also been afforded legal protection and conservation status.
There is a rapidly expanding long-line fishing industry directed at the Patagonian toothfish (Dissostichus eleginoides) in the Southern Ocean, which is marketed as Chilean sea bass. This fish, for which there is an extremely lucrative market, was only discovered and named in the early 1990s. It has a wide range, which covers a number of political zones. Since 1996, there has been widespread illegal fishing of this resource. Many fishing boats fly flags of convenience or are unflagged. Heavily implicated in this illegal fishery are Chilean operators, in which country there exists a highly developed illegal network to process the fish products.
Concentrated numbers of Dissostichus eleginoides are found around sub-Antarctic islands, including Heard and the Prince Edward, Crozet, Kerguelen Islands within the ecoregion. The level of fishing taking place is threatening the sustainability of the resource. At Heard Island, it has been suggested that the D. eleginoides fishery will most likely affect the southern elephant seal (Mirounga leonina), which has already seen a dramatic population decrease, possibly due to overfishing around the Kerguelen Islands. Overfishing has been linked to M. leonina population declines elsewhere in the ecoregion, although it should be mentioned again here that the latest studies suggest that Mirounga leonina populations are stabilizing at all sites.
Biologically significant numbers of birds are killed by hooks laid by long-line fisheries. This is especially true of uncontrolled fisheries such as this, where mitigation measures (see below) are obviously not in place. Particularly affected are albatrosses. Diomedea exulans populations at Crozet and Kerguelen have showed a slow increase since 1986, since the Japanese fishing effort for the southern blue tuna (Thunnus maccoyii) was decreased in this region. However, given that younger and more naïve birds would be more affected, there should be a time lag of 5-10 years before a decrease in the wandering albatross’ breeding population is reflected. This suggests that perhaps the consequences of the recently developed intense Dissostichus eleginoides fishery must still be fully reflected by breeding populations of Diomedea exulans. There is some evidence that foraging wandering albatrosses follow the known movements of long-line fishing fleets.
Diving species such as the gray-headed albatross (Thalassarche chrysostoma) and particularly the white-chinned petrel (Procellaria aequinoctialis) are very vulnerable to the fishery. Certainly, P. aequinoctialis forms the bulk of the avian by-catch in long-line fisheries. P. aequinoctialis already has a strong natural tendency to follow Orcinus orca pods in order to scavenge from their kills. The black-browed albatross (Thalassarche melanophrys) also does this, but to a lesser extent. Such natural habits make these pelagic birds more vulnerable to humans mimicking O. orca hunting activity. Studies on Crozet Island have shown that P. aequinoctialis forages widely, and that, during incubation, its average foraging range is the longest recorded for any seabird, foraging well beyond the Southern Ocean. This makes P. aequinoctialis, along with other birds that forage widely, particularly vulnerable to fishing activity outside protected areas. P. aequinoctialis can also forage at night, reducing the effectiveness of nighttime fishing, which has been used in the legal fishery to reduce fishery-induced albatross mortality.
The population declines of Thalassarche melanophrys at the Kerguelen Islands have also been linked to fishing activity, and it too has been recorded as a ship-following species. The fisheries are also a suggested cause for the decline in Phoebetria fusca and Thalassarche carteri populations. Apart from these three albatrosses, it is suspected that the giant petrels (Macronectes spp.) are also being negatively affected by the fishery.
Pollution: Potentially harmful releases of Polychlorinated Biphenyls (PCBs) into the atmosphere through the burning of plastic waste by island researchers, potential leaks from fuel tanks on island stations, entanglement by and ingestion of fishing debris by the islands’ fauna, and oil pollution (in 1992 about 200 oiled Aptenodytes patagonicus penguins were found on the Crozet archipelago, and in 1980 a Soviet tanker spilt 600 tonnes of petroleum after running aground north of Grande Terre).
Disturbance of King Penguin colonies: Aptenodytes patagonicus are extremely susceptible to noise stress, even showing extreme reactions to thunderstorms. Human-induced examples of this include building activities that took place in the middle of a colony of A. patagonicus on Île de la Possession of the Crozet group, which caused the breeding birds to move elsewhere. It has been suggested that the many King Penguins found dead in 1990 on Macquarie Island (outside the ecoregion) after an aircraft had flown by was caused by panic. Similar panic, although no recorded deaths, has been observed on Marion Island.
Other human activity: Collection of artifacts by island-based personnel and research huts, the latter serving as focal points for the introduction and spread of alien biota.
Global warming: Smith has warned that the effects of climate change on Marion Island could result in opportunities for new organisms to colonize, and affect the breeding success of birds. The same is true of the other sub-Antarctic islands. Also possible is the release of current environmental constraints on already introduced aliens (e.g., Mus musculus on Marion Island). It has also been shown that the breeding performances of some seals and birds in the ecoregion are reduced by El Niño Southern Oscillation (ENSO) events.
- Arnaud P. M. 1972. Ecological notes on Zanclorhynchus spinifer (Teleosti, Congiopodidae) at the Kerguelen Islands and on its predation by Giant petrels Macronectus spp. Tethys 4(3):757-760.
- Bartsch I. 1999. Peregrinacarus reticulatus gen. nov. spec. nov., a freshwater halacarid mite (Acari, Halacaridae) from Marion Island. Hydrobiologia. 392:225-232.
- Bateman S. and Rothwell D. R. (editors). 1998. Southern ocean fishing: Policy challenges for Australia. Wollongong Pap. Marit. Policy 7:140 pp.
- Bellair-Roche N. 1972. Palynological study of a bog in Vallee des Branloires, Ile de la Possession, Iles Crozet. In: Adie R. J. (ed.) Antarctic geology and geophysics. Universitetsforlaget. Oslo:831-834. ISBN: 8200022536
- Benninghoff W. S. 1987. The Antarctic ecosystem. Environment International 13:9-14.
- Benon P., Murail J. F. and Arnaud P. M. 1979. Hydrology of the peri-insular platforms of Crozet, Marion and Prince Edward Islands. (Hydrologie des Plateformes Peri-Insulaires des Iles Crozet, Marion et Prince Edward.). C. N. F. R. A. 44:39-60.
- Berruti A. 1979. The breeding biologies of the sooty albatrosses Phoebetria fusca and P. palpebrata. Emu. 79(4):161-175.
- Bester M. N. 1984. Status of the populations of the fur seals Arctocephalis tropicalis and Arctocephalis gazella north of the Antarctic Convergence. South African Journal of Science 80:27-28.
- Bester M. N. and Bartlett P. A. 1990. Attendance behaviour of Antarctic and Subantarctic fur seal females at Marion Island. Antarctic Science 2(4):309-312.
- Bester M. N. and Buchner H. 2000. Final eradication of feral cats from sub-Antarctic Marion Island, southern Indian Ocean. South African Journal of Wildlife Research 30(1):53-57.
- Bester M. N. and Miller D. G. M. 1987. Southern elephant seals and CCAMLR. Selected Scientific Papers CCAMLR 1987:465-473.
- Bester, M. N. and Pansegrouw, H. M. 1992. Ranging behaviour southern elephant seal cows from Marion Island. South African Journal of Science 88(11-12):574-575.
- Bester M. N. and van Jaarsveld A. S. 1994. Sex-specific and latitudinal variance in postnatal growth of the Subantarctic fur seal (Arctocephalus tropicalis). Canadian Journal of Zoology 72(6):1126-1133.
- Blankley W. O. and Branch G. M. 1985. Ecology of the limpet Nacella delesserti (Philippi) at Marion Island in the sub-antarctic Southern Ocean. Journal of Experimental Marine Biology and Ecology 92(2-3):259-281.
- Blankley W. O. and Grindley J. R. 1985. The intertidal and shallow subtidal food web at Marion Island. In: Siegfried, W. R., Condy P. R. and Laws, R. M. (eds.). Antarctic nutrient cycles and food webs. Springer-Verlag. New York: 630-636. ISBN: 0387134174
- Bloomer J. P. and Bester M. N. 1990. Diet of a declining feral cat Felis catus population on Marion Island. South African Journal of Wildlife Research 20(1):1-4.
- Bonner W. N. and Walton D. W. H. 1985. Key environments - Antarctica. Pegramon Press. Oxford. ISBN: 0080288812
- Borsa P. 1990. Seasonal occurrence of the leopard seal Hydrurga leptonyx in the Kerguelen Islands. Canadian Journal of Zoology 68(2):405-408.
- Branch M. L., Griffiths C. L., Kensley B. and Sieg J. 1991. The benthic Crustacea of Subantarctic Marion and Prince Edward Islands: illustrated keys to the species and results of the 1982-1989 University of Cape Town surveys. South African Journal of Antarctic Research 21(1):3-44.
- Bried J. and Jouventin P. 1997. Morphological and vocal variation among subspecies of the Black-faced Sheathbill. Condor 99(3):818-825.
- Burger A. E. 1978. Interspecific breeding attempts by Macronectus giganteus and M. halli. Emu 78:234-235.
- Burke D. 1998. Fish and seabirds threatened in the southern oceans. Antarctic 16(1):4-21.
- Cardenas J. C. and Melillanca P. I. 1999. The Chilean fishing industry: Its involvement in and connections to the illegal, unreported and unregulated exploitation of Patagonian toothfish in the Southern Ocean. Isofish Occas. Rep. 2:98pp.
- Chapuis J. L., Bousses P. and Barnaud G. 1994. Alien mammals, impact and management in the French Subantarctic islands. Biological Conservation 67:97-104.
- Chekchak T., Chapuis J. L., Pisanu B. and Bousses P. 2000. Introduction of the rabbit flea, Spilopsyllus cuniculi (Dale), to a Subantarctic island (Kerguelen Archipelago) and its assessment as a vector of myxomatosis. Wildlife Research. 27(1):91-101.
- Cherel Y., Verdon C. and Ridoux V. 1993. Seasonal importance of oceanic myctophids in king penguin diet at Crozet Islands. Polar Biology 13(5):355-357.
- Cherel Y., Weimerskirch H. and Duhamel G. 1996. Interactions between longline vessels and seabirds in Kerguelen waters and a method to reduce seabird mortality. Biological Conservation 75(1):63-70.
- Chown S. L. 1990. Possible effects of Quarternary climatic change on the composition of insect communities of the South Indian Ocean Province Islands. South African Journal of Science 86:386-391.
- Chown S. L. and Language K. 1994. Recently established Diptera and Lepidoptera on sub-Antarctic Marion Island. African Entomology 2(1):57-60.
- Clark M. R. and Dingwall P. R. 1985. Conservation of islands in the Southern Ocean: a review of the protected areas of Insulantarctica. IUCN. Cambridge.
- Cooper J. 1995a. Conservation of the Prince Edward islands. In: Dingwall P. R. (ed). Progress in conservation of the Subantarctic islands. IUCN. Cambridge: 21-30. ISBN: 2831702577
- Cooper J. 1995b. Introduced island biota: discussion and recommendations. In: Dingwall P.R. (ed). Progress in conservation of the Subantarctic islands. IUCN. Cambridge: 133-138. ISBN: 2831702577
- Cooper J. and Brown C. R. 1990. Ornithological research at the sub-Antarctic Prince Edward Islands: a review of achievements. South African Journal of Antarctic Research 20(2):40-57.
- Cooper J., Crafford J. E. and Hecht T. 1992. Introduction and extinction of brown trout (Salmo trutta L.) in an impoverished Subantarctic stream. Antarctic Science 4(1):9-14.
- Cooper J., Avenant N. L. and Lafite P. W. 1994. Airdrops and king penguins: A potential conservation problem at sub-Antarctic Marion Island. Polar Record. 30(175):277-282.
- Cooper J., Marais A. v.N., Bloomer J. P. and Bester M. N. 1995. A success story: Breeding of burrowing petrels (Procellariidae) before and after the eradication of feral cats Felis catus at Subantarctic Marion Island. Marine Ornithology 23(1):33-37.
- Cox J. B. 1978. Albatross killed by giant-petrel. Emu 78(2):94-95.
- Crafford J. E. and Chown S. L. 1987. Plutella xylostella L. (Lepidoptera: Plutellidae) on Marion Island. Journal of the Entomological Society of Southern Africa 50(1):259-260.
- Croxall J. P. and Gales R. 1997. An assessment of the conservation status of albatrosses. In: Robertson G. and Gales R. (eds.). Albatross biology and conservation. Surrey Beatty and Sons. Chipping Norton: 46-65. ISBN: 0949324825
- Dalziell, J. and Poorter, M. 1993. Seabird mortality in longline fisheries around South Georgia. Polar Record 29(169):143-145.
- Davaine P. 1991. Sea ranching of coho salmon in the Kerguelen Islands (TAAF). ICES: Copenhagen. 21pp.
- Davaine P. and Beall E. 1982. Introductions of salmonids in the French Austral Territories. C. N. F. R. A. 51:289-300.
- DEAandT. 1996. Department of Environmental Affairs and Tourism. Prince Edward Islands Management Plan. DEAandT. Pretoria.
- Deacon G. E. R. 1983. Kerguelen, Antarctic and Subantarctic. Deep-Sea Res. 30(1A):77-81.
- Dingwall P. R. 1995. Progress in conservation of the Subantarctic islands. IUCN. Gland and Cambridge. 225 pp. ISBN: 2831702577
- du Plessis C. J., van Heezik Y. M. and Seddon P. J. 1994. Timing of king penguin breeding at Marion Island. Emu 94(3):216-219.
- Duhamel G. 1998. The pelagic fish community of the Polar frontal zone of the Kerguelen Islands. In: Di Prisco G., Pisano E. and Clarke A. (eds.). Fishes of Antarctica. A biological overview. Springer-Verlag Italia. Milano: 63-74. ISBN: 8847000289
- Frenot Y., Vernon P. and Bellido A.. 1989. A bibliography of terrestrial ecosystems on Iles Crozet, Indian Ocean. Polar Record 25(153):121-130.
- Frey F. A., Weis D., Yang H. J., Nicolaysen K., Leyrit H. and Giret A. 2000. Temporal geochemical trends in Kerguelen Archipelago basalts; evidence for decreasing magma supply from the Kerguelen Plume. Chemical Geology 164(1-2):61-80.
- Froneman P. W. and Ansorge I. J. 1998. The Third Marion Island Oceanographic Study (MIOS III) conducted during April and May 1998. South African Journal of Science 94(9):437-439.
- Gales R. 1993. Cooperative mechanisms for the conservation of albatross. Australian Antarctic Foundation. Hobart.
- Gauthier C. M., Clerquin Y. and Drault S. 1998. Desertion of a King Penguin colony (Aptenodytes patagonicus) following a thunderstorm Alauda 66(3):195-198.
- Girod M. and Nougier J. 1972. Volcanism of the sub-Antarctic islands. In: Adie R.J. (ed.) Antarctic geology and geophysics. Universitetsforlaget. Oslo: 777-788. ISBN: 8200022536
- Goldsworthy S. D. and Shaughnessy P. D. 1989. Subantarctic fur seals Arctocephalus tropicalis at Heard Island. Polar Biology 9(5):337-339.
- Green K. and Burton H. R. 1993. Comparison of the stomach contents of southern elephant seals, Mirounga leonina, at Macquarie and Heard Islands. Marine Mammal Science 9(1):10-22.
- Greene S. W. and Walton D. W. H. 1975. An annotated check list of the sub-antarctic and antarctic vascular flora. Polar Record 17(110):473-484.
- Green K., Slip D. J. and Moore G. J. 1998. The take of fish species by seabirds and marine mammals in the Australian Fisheries Zone around Heard Island: The potential for competition with a commercial fishery. Polar Biology 20(4):273-280.
- Gregoire M., Mattielli N., Nicollet C., Cottin J. Y., Leyrit H., Weis D., Shimizu N. and Giret A. 1994. Oceanic mafic granulite xenoliths from the Kerguelen archipelago. Nature 367:360-363.
- Gremmen N. J. M. 1981. The vegetation of the Subantarctic islands Marion and Prince Edward. Dr W. Junk. The Hague. 149pp.
- Gremmen N. J. M., Chown S. L. and Marshall D. J. 1998. Impact of the introduced grass Agrostis stolonifera on vegetation and soil fauna communities at Marion Island, sub-Antarctic. Biological Conservation 85(3):223-231.
- Gribnitz K. H., Kent L. E. and Dixon R. D. 1986. Volcanic ash, ash soils and the inferred Quaternary climate of sub-Antarctic Marion Island. South African Journal of Science 82(11):629-635.
- Grindley J. R., David P. and Condy P. R. 1985. Nutrient upwelling and its effects in the lee of Marion Island. In: Siegfried, W. R., Condy P. R. and Laws, R. M. (eds.). Antarctic nutrient cycles and food webs. Springer-Verlag. New York: 46-51. ISBN: 0387134174
- Grobbelaar J. U. 1978. The limnology of Marion Island: southern Indian Ocean. South African Journal of Antarctic Research 8:113-118.
- Guinet, C. 1991. Intentional stranding apprenticeship and social play in killer whales (Orcinus orca) Canadian Journal of Zoology 69(11):2712-2716.
- Guinet C. and Bouvier J. 1995. Development of intentional stranding hunting techniques in killer whale (Orcinus orca) calves at Crozet Archipelago. Canadian Journal of Zoology 73(1):27-33.
- Guinet C., Jouventin P. and Weimerskirch H. 1992. Population changes, movements of southern elephant seals on Crozet and Kerguelen archipelagos in the last decades. Polar Biology 12(3-4):349-356.
- Guinet C., Jouventin P. and Georges J. Y. 1994. Long-term population changes of fur seals Arctocephalus gazella and Arctocephalus tropicalis on Subantarctic (Crozet) and subtropical (St. Paul and Amsterdam) islands and their possible relationship to El Niño Southern Oscillation. Antarctic Science 6(4):473-478.
- Guinet C., Chastel O., Koudil M., Durbec J. P. and Jouventin P. 1998. Effects of warm sea-surface temperature anomalies on the blue petrel at the Kerguelen Islands. Proceedings of the Royal Society of London, Series B: Biological Sciences 265(1400):1001-1006.
- Guinet C., Jouventin P. and Weimerskirch H. 1999. Recent population change of the southern elephant seal at Iles Crozet and Iles Kerguelen: the end of the decrease? Antarctic Science 11(2):193-197.
- Hänel C. and Chown S. 1998. An introductory guide to the Marion and Prince Edward Island Special Nature Reserves. DEAandT. Pretoria.
- Hay C. H. 1986. A new species of Macrocystis C. Ag. (Phaeophyta) from Marion Island, southern Indian Ocean. Phycologia 25(2):241-252.
- Higgins P. J. and Davies S. J. J. F. (editors) 1996. Handbook of Australian, New Zealand and Antarctic birds. Vol. 3. Oxford University Press. Melbourne. ISBN: 0195530705
- Hofmeyr G. J. G., Bester M. N. and Jonker, F. C. 1997. Changes in population sizes and distribution of fur seals at Marion Island. Polar Biology 17(2):150-158.
- Hughes J. M. R. 1987. The distribution and composition of vascular plant communities on Heard Island. Polar Biology 7:153-162.
- Hunter S. 1985. The role of giant petrels in the Southern Ocean ecosystem. In: Siegfried W. R., Condy P. R. and Laws R. M. (eds.) Antarctic nutrient cycles and food webs. Springer-Verlag. New York: 534-542. ISBN: 0387134174
- Hunter S. 1991. The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island. Ibis 133(4):343-350.
- Hunter S. and Brooke M. 1992. The diet of giant petrels Macronectes spp. at Marion Island, Southern Indian Ocean. Colonial Waterbirds 15(1):56-65.
- Huyser O., Ryan P. G. and Cooper J. 2000. Changes in population size, habitat use and breeding biology of lesser sheathbills (Chionis minor) at Marion Island: Impacts of cats, mice and climate change? Biological Conservation 92(3):299-310.
- IUCN 2000. 2000 IUCN red list of threatened species.
- Johnstone G. W. 1978. Interbreeding by Macronectus halli and M. giganteus at Macquarie Island. Emu 78:235.
- Johnstone G. W., Shaughnessy P. D. and Conroy J. W. H. 1976. Giant-petrels in the South Atlantic: new data from Gough Island. South African Journal of Antarctic Research 6:19-22.
- Jouventin P. and Micol T. 1995. Conservation status of the French Subantarctic Islands. In: Dingwall P. R. (ed.) Progress in conservation of Subantarctic islands. IUCN. Cambridge: 31-42. ISBN: 2831702577
- Jouventin P., Bried J. and Ausilio E. 1996. Life-history variations of the Lesser Sheathbill Chionis minor in contrasting habitats. Ibis 138(4):732-741.
- Kerley G. I. H. 1984. Relationships between sympatric breeding populations of fur seals (Arctocephalis spp.) at the Prince Edward Islands. South African Journal of Science 80:28-30.
- Kerley G. I. H. 1989. Electrophoretic transferrin variation in fur seals (Arctocephalus spp.) at Marion Island. Comp. Biochem. Physiol. B. 92B(2):361-364.
- Kessler J. 1999. HEADLINE: A fish in hot water; Chilean sea bass is increasing in price, decreasing in supply and may be tailing off the menu. The Atlanta Journal and Constitution. March 26, 1999, Friday, CONSTITUTION EDITION.
- Klyausov A. V. 1994. Rapprochement and confluence of climatic fronts eastward of the Kerguelen Archipelago. Ecological effects. Tr. YugNIRO/Proc. South. Sci. Res. Inst. Mar. Fish. Ocean. 40:26-30.
- Knox G. A. 1977. The Antarctic polychate fauna: its characteristics, distribution patterns and evolution. In: Llano G. A. (ed.). Adaptations within Antarctic ecosystems - Proceedings of the third SCAR symposium on Antarctic biology. Gulf Publishing Company. Houston. ISBN: 0872010007
- Koubbi B. 1993. Influence of the frontal zones on ichthyoplankton and mesopelagic fish assemblages in the Crozet Basin (Indian sector of the Southern Ocean). Polar Biology 13(8):557-564.
- Lane M. 1998. HEADLINE: NZ crackdown on Ross Sea fishing pirates The Evening Post (Wellington). December 29, 1998.
- Lewis-Smith R. I. 1984. Terrestrial plant biology. In: Laws R. M. (ed.) Antarctic Ecology. Vol. 1. Academic Press. London: 61-162. ISBN: 0124395023
- Lugten G. L. 1997. The rise and fall of the Patagonian toothfish - food for thought. Environmental Policy and Law 27(5):401-407.
- Marchant S. and Higgins P. J. (co-ordinators). 1990. Handbook of Australian, New Zealand and Antarctic birds. Vol. 1 (AandB). Oxford University Press. Melbourne. ISBN: 0195530683
- Marchant S. and Higgins P.J. (editors). 1993. Handbook of Australian, New Zealand and Antarctic birds. Vol. 2. Oxford University Press. Mebourne. ISBN: 0195530691
- Merten M. 2000. Troubled waters for SA's 'white gold'. Daily Mail and Guardian. December 27, 2000.
- Michalev J. A. 1978. Revealing of differences in the Antarctic baleen whale stocks on the basis of the analysis of the ‘Jacobson's organ’ position. Rep. Int. Whaling Comm. In: Twenty-ninth report of the International Whaling Commission covering the twenty-ninth fiscal year 1977-1978. IWC; Cambridge: 343-346.
- Moloney C. L., Cooper J., Ryan P. G. and Siegfried W. R. 1994. Use of a population model to assess the impact of longline fishing on wandering albatross Diomedea exulans populations. Biological Conservation 70(3):195-203.
- Newton I. P. and Fugler S. R. 1989. Notes on the winter-breeding Greatwinged petrel Pterodroma macroptera and Gray petrel Procellaria cinerea at Marion Island. Cormorant 17(1-2):27-34.
- Nicolaysen K., Frey F. A., Hodges K. V., Weis D. and Giret A. 2000. 40AR139AR geochronology of flood basalts from the Kerguelen Archipelago, southern Indian Ocean: implications for Cenozoic eruption rates of the Kerguelen Plume. Earth and Planetary Science Letters 174(3-4):313-328.
- Park, Y. H., Gamberoni L. and Charriaud E. 1993. Frontal structure, water masses, and circulation in the Crozet Basin. Journal of Geophysical Research 98(C7):12 361-12 385.
- Perissinotto R. and McQuaid C. 1990. Role of the sub-antarctic shrimp Nauticaris marionis in coupling bethic and pelagic food webs. Marine Ecology Progress Series 64:81-87.
- Perissinotto R. and McQuaid C. D. 1992. Deep occurrence of the giant kelp Macrocystis laevis in the Southern Ocean. Marine Ecology Progress Series 81(1):89-95.
- Perrin R. A. 1995. Conservation status of Heard Island and McDonald Islands. In: Dingwall P. R. (ed.) Progress in conservation of the Subantarctic islands. IUCN. Cambridge: 43-52. ISBN: 2831702577
- Prince P. A., Weimerskirch H., Huin N. and Rodwell S. 1997. Molt, maturation of plumage and ageing in the wandering albatross. Condor 99(1):58-72.
- Punta G. and Herrera G. 1995. Predation by southern giant petrels Macronectes giganteus on adult imperial cormorants Phalacrocorax atriceps. Marine Ornithology 23(2):166.
- Reale D., Bousses P., Pisanu B. and Chapuis J. L. 2000. Biannual reproductive cycle in the Kerguelen feral sheep population. Journal of Mammalogy 81(1):169-178.
- Recq M., Goslin J., Charvis P. and Operto S. O. 1998. Small-scale crustal variability within an intraplate structure; the Crozet Bank (southern Indian Ocean). Geophysical Journal International 134(1):145-156.
- Reilly A. and Ward R. D. 1999 Microsatellite loci to determine population structure of the Patagonian toothfish Dissostichus eleginoides. Molecular Ecology 8(10):1753-1754.
- Ridoux V. 1987. Feeding association between seabirds and killer whales, Orcinus orca, around Subantarctic Crozet Islands. Canadian Journal of Zoology 65(8):2113-2115.
- Ridoux V. 1994. The diets and dietary segregation of seabirds at the Subantarctic Crozet Islands. Marine Ornithology 22(1):1-192.
- Ryan P. G. and Boix-Hinzen C. 1998. Tuna longline fisheries off southern Africa: the need to limit seabird bycatch. South African Journal of Science 94(4):179-182.
- Saunders S. 2000. Catches of Patagonian toothfish to be tracked. South African Shipping News and Fishing Industry Review 54(1):17.
- Slip D. J. and Burton H. R. 1999. Population status and seasonal haulout patterns of the southern elephant seal (Mirounga leonina) at Heard Island. Antarctic Science 11(1):38-47.
- Smith V. R. 1985. Heterotrophic acetylene reduction in soils at Marion Island. In: Siegfried W. R., Condy P. R. and Laws R. M. (eds.) Antarctic nutrient cycles and food webs. Springer-Verlag. New York: 186-191. ISBN: 0387134174
- Smith V. R. 1987. The environment and biota of Marion Island. South African Journal of Science 83(4):211-220.
- Smith V. R. 1991. Climate change and its ecological consequences at Marion and Prince Edward Islands. South African Journal of Antarctic Research 21(2):223-224.
- Smith V. R. and Lewis Smith R. I. 1987. The biota and conservation status of sub-Antarctic islands. Environment International 13:95-104.
- Smith V. R. and Steenkamp M. 1992. Macroinvertebrates and litter nutrient release on a sub-Antarctic Island. South African Journal of Botany 58(2):105-116.
- Steenkamp W. 2000. Fully fledged coastguard to fight poachers. Saturday Argus. December 9/10 2000.
- Tuck G. N., Polacheck T., Croxall J. P., Weimerskirch H., Prince P. A., and Wotherspoon S. 1999. The potential of archival tags to provide long-term movement and behaviour data for seabirds: first results from wandering albatross Diomedea exulans of South Georgia and the Crozet Islands. Emu 99(1):60-68.
- Udvardy M. D. F. 1975. A classification of the biogeographical provinces of the world. IUCN Occasional Paper 18.
- van Rensburg P. J. J. and Bester M. N. 1988. The effect of cat Felis catus predation on three breeding Procellariidae species on Marion Island. South African Journal of Zoology 23(4):301-305.
- van Tussenbroek B. I. 1989. Smooth-bladed Macrocystis (Laminariales, Phaeophyta) from the Falkland Islands. Phycologia 28(4):520-523.
- van Zinderen Bakker E. M. (Snr.), Winterbottom J. M. and Dyer R. A. 1971. Marion and Prince Edward islands: Report on the South African Biological and Geological expedition 1965-1966. AA Balkema. Cape Town.
- Verwoerd W. J., Russell S. and Berruti A. 1981. 1980 Volcanic eruption reported on Marion Island. Earth and Planetary Science Letters 54(1):153-156.
- WCMC 1997. World Conservation Monitoring Centre. Descriptions of natural World Heritage Properties.
- Weimerskirch H. and Stahl J. C. 1988. The breeding and feeding ecology of the Kerguelen tern Sterna virgata. Ornis Scandinavica 19(3):199-204.
- Weimerskirch H., Jouventin P. and Stahl J. C. 1986. Comparative ecology of the six albatross species breeding on the Crozet Islands. Ibis 128(2):195-213.
- Weimerskirch H., Zotier R. and Jouventin P. 1988. The avifauna of the Kerguelen Islands. Emu 89:15-29.
- Weimerskirch H., Brothers N. and Jouventin P. 1997. Population dynamics of wandering albatross Diomedea exulans and Amsterdam albatross D. amsterdamensis in the Indian Ocean and their relationships with long-line fisheries: Conservation implications. Biological Conservation 79(2-3):257-270.
- Weimerskirch H., Catard A., Prince P. A., Cherel Y. and Croxall J. P. 1999. Foraging white-chinned petrels Procellaria aequinoctialis at risk: from the tropics to Antarctica. Biological Conservation 87(2):273-275.
- Williams A. J. 1980. Aspects of the breeding biology of the Gentoo Penguin, Pygoscelis papua. Gerfaut/Giervalk 70(3):283-295.
- Williams A. J., Siegfried W. R., Burger A. E. and Berruti A. 1979. The Prince Edward islands: a sanctuary for seabirds in the Southern Ocean. Biological Conservation. 15(1):59-71.
- Winterbottom J. M. 1971. The position of Marion Island in the Sub-Antarctic avifauna. In: van Zinderen Bakker E. M. (snr.), Winterbottom J. M. and Dyer R. A. (eds.). Marion and Prince Edward Islands: Report on the South African Biological and Geological expedition 1965-1966. Balkema. Cape Town: 241-248.
- Woehler E. J. 1991. Morphology of prions Pachyptila and diving petrels Pelecanoides at Heard Island. Marine Ornithology 19(1):19-30.
- Woehler E. J. and Gilbert C. A. 1990. Hybrid rockhopper-macaroni penguins, interbreeding and mixed species pairs at Heard and Marion islands. Emu 90(3):198-210.
- Yau, C., Collins M. A., Everson I. and Nolan C. P. 1998. A new method for estimating the abundance of Patagonian toothfish (Dissostichus eleginoides). ICES Copenhagen. 13 pp.
- Zdzitowiecki K. and Pisano E. 1996. New records of acanthocephalans infecting fish off Heard Island (Kerguelen sub-region, sub-Antarctic). Archive of fishery and marine research 43(3):257-264.
- Zhivov V. V. and Krivoruchko V. M. 1990. On the biology of the patagonian toothfish, Dissostichus eleginoides, of the Antarctic part of the Atlantic. Journal of Ichthyology 30(7):142-146.