Notothenioidei

(Southern cod-icefishes)

Class Actinopterygii

Order Perciformes

Suborder Notothenioidei

Number of families 8

Evolution and systematics

Over the past 40 million years, the suborder Notothenioidei has evolved in high latitudes of the Southern Hemisphere from generalized blennioid ancestors, producing a remarkable variety of ecological, morphological, physiological, and biochemical specializations. This diversity has led to their recognition as the only known example of a marine species swarm, rivaling the freshwater swarms of cichlid fishes in the African rift lakes and the landlocked cottoids of Lake Baikal. Over the past three decades, intensive study of the group has been driven by the increase in internationally funded research in Antarctica that followed the Antarctic Treaty of 1959.

The basic taxonomic framework for the suborder, established in the early 1900s, recognized five families; as of 2002, the Notothenioidei included 122 species in 43 genera, divided into eight families:

• Bovichtidae (thornfishes, 10 species in two genera); the most primitive family
• Pseudaphritidae (one catadromous species, Pseudaphritis urvillii, the Australian congolli or tupong)
• Eleginopidae (one species, Eleginops maclovinus)
• Harpagiferidae (spiny plunderfishes, six species in one genus)
• Artedidraconidae (plunderfishes, 25 species in four genera)
• Nototheniidae (notothens, or "Antarctic cods," 49 species in 12 genera)
• Bathydraconidae (dragonfishes; 15 species in 11 genera)
• Channichthyidae (icefishes, 15 species in 11 genera)

There are no unequivocal notothenioid fossils; thus, little is known of the origins of the suborder. Earlier claims of Eocene and Miocene notothenioid fossils from the Antarctic Peninsula and New Zealand have been discounted. The only substantial Antarctic fish fossils are found in Eocene deposits on Seymour Island, near the Antarctic Peninsula. These beds contain a cosmopolitan shallow-water fossil fish fauna that is completely different from the modern Antarctic fauna, presently dominated by notothenioids. As Antarctica became glaciated, the hardy notothenioids replaced the earlier fishes and then diversified under the impetus of habitat destruction associated with cyclical glacial advances.

Physical characteristics

Notothenioids have a diversity of sizes and body forms. Most are small fishes, on the order of 1 ft (30 cm) in length, but some species may be as small as 4.3 in (11 cm) or as long as 6 ft (1.8 m). None is highly colored; most have black, brown, or gray mottling on paler backgrounds. They have two to three lateral lines on the trunk and well-developed sensory canals on the head, jaw, and pre-operculum. Three plate-like radials characterize the pectoral girdle. All notothenioids lack a swim bladder.

Many of the 96 species of Antarctic notothenioids coexist with sea ice, near the freezing point of seawater (28.6°F, or −1.87°C). They are protected from freezing by an antifreeze glycopeptide (AFGP), consisting of repeating groups of three amino acids (-alanine-alanine-threonine-) n, with a disaccharide (galactose-N-acetylgalactosamine) attached to each threonine. AFGP works differently from normal colligative antifreezes such as salt or glycol, which only depend on the number of ions or molecules to lower freezing temperature. Instead, AFGP molecules bind to mircroscopic ice crystals and interfere with the attachement of additional water molecules, preventing ice from growing to a size that would damage living cells. Thus a very small number of AFPG molecules can have a disproportionate effect on freezing. The gene encoding AFGP has evolved from part of a gene for the digestive enzyme trypsin.

As well as being important for survival, AFGPs are metabolically expensive. In the high-latitude notothenioids that produce AFGPs, changes in the kidney reduce their loss. Instead of forming urine by pressure filtration in capillary bundles (glomeruli), Antarctic notothenioids eliminate wastes by selective secretion into the urine, and AFGPs are retained. Their kidneys lack the glomeruli characteristic of most other vertebrates. In contrast, the 26 species of notothenioids living outside Antarctic waters rarely encounter freezing seawater, lack AFGPs, and have glomeruli.

Distribution

Three of the eight notothenioid families are primarily non-Antarctic:

• Bovichtidae: These are found mainly north of the Antarctic convergence, in Australia/Tasmania, southern New Zealand, and southern South America. One species occurs along the Antarctic Peninsula.
• Pseudaphritidae: The family inhabits streams and estuaries of southeastern Australia and Tasmania.
• Eleginopidae: These fishes are confined to southern South America.

The remaining five families have distributions that are primarily Antarctic or subantarctic:

• Harpagiferidae: The family is found mainly in sub-Antarctic regions, from Patagonia, the Falkland Islands, South Georgia, and the Scotia Arc to the tip of the Antarctic Peninsula and in Kerguelen, Crozet, and Heard Islands.
• Artedidraconidae: These fishes are exclusively Antarctic, from South Georgia along the Scotia Arc to the Antarctic Peninsula and around continental Antarctica.
• Nototheniidae: Thirty-four species are truly Antarctic, and the remainder occur in sub-Antarctica, Patagonia, and southern New Zealand.
• Bathydraconidae: The family occurs in the Antarctic, from South Georgia along the Scotia Arc and around the Antarctic continent.
• Channichthyidae: The family is confined primarily to the Antarctic but has one species in Patagonia/Falkland Islands. The other 14 species are found from South Georgia through the Scotia Arc and around the Antarctic continent; they also occur in the Bouvet and Kerguelen Islands but not as far as the Heard, Campbell, or Auckland Islands.

Habitat

Notothenioids are widespread in coastal waters of the Southern Ocean, on or over the continental shelf. Their depth range is from the surface to 1,370 fathoms (2,500 m). Aside from their dispersive larval stages, most avoid the open ocean, living on or near the bottom as adults. Several species of Nototheniidae and Channichthyidae have abandoned their ancestral benthic existence, however, to move up into the water column, becoming pelagic. Lacking swim bladders, these mid-water forms have increased buoyancy in other ways. Their bones are less mineralized than those of demersal species, and they have high oil contents. One widespread pelagic species, the sardine-like nototheniid Pleuragramma antarcticum, retains larval features, including a persistent notochord surrounded by greatly reduced vertebral centra.

Behavior

Notothenioids tend to be sedentary bottom species that swim infrequently. Where swimming has been observed, it is labriform, propelled by undulatory sculling of the pectoral fins. The trunk and tail are used only for short, fast dashes. Even the pelagic forms are relatively inactive.

Feeding ecology and diet

Demersal notothenioids are generalized predators. Some forage nonselectively; others are ambush predators. Pelagic species, such as P. antarcticum, are more selective, feeding on copepods, larger midwater crustacea (e.g., krill, such as Euphausia superba), and pteropods (e.g., Limacina). In turn, these smaller midwater fishes are eaten, along with krill and squid, by the few large mesopelagic fish (such as the toothfish, Dissostichus mawsoni), by birds (penguins, skuas, and petrels), and seals. D. mawsoni is eaten by Weddell seals (Leptonychotes weddelli) and by killer whales (Orcinus orca).

Reproductive biology

Many Antarctic and subantarctic notothenioids breed biennially, with oocytes taking two years to ripen. Eggs are large and yolky and usually are spawned on or near the bottom. Spawning times differ with latitude and between species, ranging from early spring to early winter. Embryonic development and growth are slow, and most larvae hatch 6–12 months after spawning. The larval stages are pelagic, settling to the bottom after feeding in the plankton for six to nine months.

Conservation status

None of the notothenoids are on the IUCN Red List. Although the majority of notothenioids are not exploited and have widespread and abundant populations, several species have been the target of fisheries. The marbled rockcod, Notothenia rossii, has been seriously overfished but is no longer targeted, and it is recovering, albeit slowly. The Patagonian toothfish, Dissostichus eleginoides, has been exploited since the 1980s and has been given the designation of data deficient in a report on the conservation status of Australian fishes complied by the Threatened Fishes Committee of the Australian Society for Fish Biology.

After depleting the South Georgia/Falkland Islands tooth-fish fishery, attention shifted to Macquarie and Kerguelen Islands and then further west, to the southern Indian Ocean. In the mid-1990s the estimated illegal catch exceeded the legal catch, but in 1999 the Commission for the Conservation of Antarctic Marine Resources implemented a strict catch-reporting regime for toothfish, reducing the 1999/2000 illegal catch to an estimated 8,418 tons (7,637 tonnes), of a total 33,660 tons (30,536 tonnes). Since 1996 there has been a limited exploratory fishery for the closely related Antarctic toothfish, D. mawsoni, in the Ross Sea, where a catch of 1,000 tons (907 tonnes) was reported in 2001. Because most notothenioids share characteristics of low fecundity and slow growth, their fisheries are highly susceptible to overfishing and should be monitored closely.

Significance to humans

Some notothenioids have been harvested for fish meal and oil (e.g., P. antarcticum), but the group is exploited mainly for human consumption. The mackerel icefish, Champsocephalus gunnari, is the basis of small, but sustainable fisheries near South Georgia and the Kerguelen Islands. There is a ready market for the two toothfish species, D. eleginoides and D. mawsoni, which are highly palatable.

Species accounts

List of Species

Sailfin plunderfish

Histiodraco velifer

family

Artedidraconidae

taxonomy

Dolloidraco velifer Regan, 1914, McMurdo Sound, Antarctica.

other common names

None known.

physical characteristics

Attains a length of up to 7.5 in (19.2 cm) (length/depth ratio, 5:1). It has a large, depressed head with a conspicuous barbel on the chin; this barbel is one of the characteristic features of the family. The first dorsal fin is tall and narrow, consisting of only three flexible spines, and is directly above the operculum. The second dorsal fin is disproportionately tall and sail-like. The second dorsal, caudal, pectoral, and pelvic fins have brown and yellow striations. The body color is a light tan, with irregular dark blotches. Scales are absent, except for parts of the two lateral lines on the trunk.

distribution

Coastal waters of Antarctica, from the Weddell Sea clockwise to the Ross Sea.

habitat

This is a bottom-dwelling fish, found at depths of 48–2,190 ft (15–667 m). It is taken occasionally by scuba divers on the mud or gravel bottoms in McMurdo Sound and Terra Nova Bay.

behavior

In the aquarium, the sailfin plunderfish sits quietly on the bottom. The barbel is extended and occasionally twitched, mimicking a small worm. Histological examination shows that the barbel is highly innervated, with many tiny capsules resembling Pacinian corpuscles (a comomn type of pressure-sensitive sense organ amongst the vertebrates). Touching the barbel with forceps elicits a feeding lunge. This behavior implies that natural feeding may target mobile foragers, such as fish and amphipods.

feeding ecology and diet

Very little is known about the diet of the sailfin plunderfish, other than what can be inferred from behavioral observations in the aquarium. Where they have been examined, gut contents include krill and polychaete worms. Like most of the benthic notothenioids, it is probably an opportunistic feeder.

reproductive biology

Nothing is known of the reproductive biology, as very few specimens have been collected.

conservation status

Although this is an uncommon species, poorly represented in museum collections, it is not threatened.

significance to humans

None known.

Naked dragonfish

Gymnodraco acuticeps

family

Bathydraconidae

taxonomy

Gymnodraco acuticeps Boulenger, 1902, Cape Adare, North Victoria Land, Antarctica.

other common names

None known.

physical characteristics

Grows to 13.5 in (34 cm) in length. It has an elongated, scaleless body (length/depth ratio, 9:0) with a long, pointed head. The coloring is yellowish to olive-brown, with darker blotches on the sides; it is pale on the ventral surface. The lower jaw extends well beyond the upper and bears two prominent fangs that are slanted backwards to prevent the escape of prey. In common with all members of the dragonfish family, it lacks the first spinous dorsal fin. This species produces abundant mucus when it is freshly caught.

distribution

South Shetland, the Antarctic Peninsula, and the eastern Antarctic continental shelf from the Weddell Sea to the Ross Sea.

habitat

Found mainly in shallow inshore waters, to a depth of 162 ft (50 m), although it has been caught as deep as 1,800 ft (550 m). It commonly resides in crevices or under rock ledges or anchor ice.

behavior

This is an aggressive fish that rapidly consumes other aquarium inhabitants.

feeding ecology and diet

The diet varies with location. Near the Antarctic Peninsula, the dragon fish feeds on krill; in McMurdo Sound, fish, amphipods, fish eggs, and polychaetes are taken.

reproductive biology

Spawning occurs in September; 0.12-in (3-mm) eggs are attached to a stone as a flattened patch of about a thousand and guarded. In the wild, adults tend egg masses until the larvae hatch and disperse. Development takes about a year. Free-swimming larvae, about 0.6 in (15 mm) long, hatch out in early summer and remain in the zooplankton for about six months.

conservation status

Not threatened.

significance to humans

This species is proving to be a valuable research subject. Beginning in 2001 clusters of developing embryos near McMurdo Station have been harvested for investigation into the ontogeny of AFGPs.

Mackerel icefish

Champsocephalus gunnari

family

Channichthyidae

taxonomy

Champsocephalus gunnari Lönnberg, 1905, Cumberland Bay, South Georgia.

other common names

English: Crocodile icefish, pike glassfish; French: Poisson des glaces; Spanish: Draco rayado; Russian: Ledyanaya ryba.

physical characteristics

Elongated, scaleless, pikelike body (length/depth ratio, about 7:2) with an elongated snout. The jaws do not protract. In coloring it is silvery gray, darker on the back, and silvery on the belly, with dark vertical stipes on the sides, reminiscent of those seen in mackerel. The gills are a pale yellowish; the blood is colorless, completely lacking hemoglobin. The absence of hemoglobin is a remarkable feature of all 15 icefish species, apparently caused by a single massive mutation that deleted the β-globin gene. Far from being an adaptation to Antarctic conditions, this has been termed a "disaptation." Several factors have enabled channichthyids to survive this evolutionary catastrophe: frigid seawater and blood plasma can carry more dissolved oxygen, and low body temperatures limit metabolic requirements. Subsequent evolution has compensated for the loss of an oxygen-binding pigment: blood volume, heart, vessel, and gill size and the perfusion rate of gills and blood vessels all have increased. As a consequence, channichthyids are surprisingly active, competing successfully with other notothenioids.

distribution

South Georgia and the islands of the Scotia Arc, southward to the northern Antarctic Peninsula; it also inhabits Bouvet, Kerguelen, and Heard Islands. A closely related species, C. esox, occurs in the Falklands Islands and Patagonia and is the only channichthyid to occur outside Antarctic/subantarctic waters.

habitat

This is a coastal species, found mainly between 330 and 1,140 ft (100–350 m). Mature adults are found offshore in summer, moving inshore to spawn in the fall (March–May).

behavior

The species aggregates in locations with dense krill populations, remaining near the bottom during the day and migrating upward with the krill at night.

feeding ecology and diet

The mackerel icefish feeds mainly on krill (Euphausia superba); it takes other euphausiids, mysids, and hyperiid amphipods (Themisto gaudichaudii) when krill numbers are low.

reproductive biology

Reproductive parameters vary considerably between the different stocks. Maturation of oocytes takes less than a year, producing 1,300–31,000 eggs that are 0.12–0.16 in (3–4 mm) in diameter. When krill are scarce, oocytes may be resorbed, and as many as 60% of mature females may be nonreproductive. A marked three-year periodicity in reproduction is reported from the Kerguelen stock. Near South Georgia, spawning occurs from March to May; it takes place later in South Orkney and South Shetland and as late as July to August in the Kerguelen stock. Eggs are deposited in the depth range of 330–660 ft (100–200 m) and hatch after 30–180 days, depending on the stock. Newly hatched larvae are 0.05–0.07 in (12–17 mm) long and feed on copepods within 12 mi (20 km) offshore; larger juvenile fish move further offshore but remain over the continental shelf. Juveniles grow at 2.5–3.5 in (6.4–8.9 cm) per year, reaching sexual maturity at three years at a length of 10 in (25 cm).

conservation status

Not listed by the IUCN. The various stocks of mackerel icefish have been targeted by minor trawl fisheries since 1974. Catches fluctuate from year to year, but they appear to be sustainable for the South Georgia and Kerguelen stocks. In South Orkney and South Shetland, the stock was depleted rapidly below sustainable levels and is no longer being fished.

significance to humans

A total catch of 4,295 tons (3,896 tonnes) was reported from the Commission for the Conservation of Antarctic Marine Resources management area in 1999–2000.

Maori chief

Notothenia [Parenotothenia] angustata

family

Nototheniidae

taxonomy

Notothenia angustata Hutton, 1875, Dunedin Harbour, New Zealand.

other common names

English: Black cod.

physical characteristics

Grows to 2 ft (60 cm) in length. Mature specimens have a massive body (length/depth ratio, 5:1) with a large, slightly flattened head; heavy bony ridges over the eyes; and large scales. The coloring is dark olive-gray to blue-black, with varying light spots and lines; there are vague dark vertical bands on the trunk. Juveniles are less flattened, with deeper bodies, deard reddish bronze coloring, and prominent light spots.

distribution

Southern waters of New Zealand, from Stewart Island to Kaikoura. It also is found in Chatham, Snares, Auckland, and Campbell Islands and in Punta Arenas, Chile, and adjacent water.

habitat

Adults are demersal on rocky bottoms, from near shore to 330 ft (100 m) depth. Juveniles are found in tide pools and among kelp near the shore.

behavior

Like most other notothenioids, the Maori chief is solitary and never occurs in schools. Adults are shy and sedentary, foraging among rocks and seaweed mainly at night. During the day, they hide in crevices or under seaweed, occasionally lunging for passing prey. Smaller fish are sucked into their mouth by the sudden expansion of the jaws and opercula. Juveniles are more active than adults, often leaving the bottom to forage in the upper layers of kelp beds.

feeding ecology and diet

The Maori chief feeds on a wide range of bottom invertebrates, crabs, and smaller fishes. Occasionally, it ingests seaweed.

reproductive biology

The life cycle of the Maori chief is not well known. Eggs ripen in November and presumably are deposited on the bottom. Juveniles appear in rock pools in the summer (January) after an incubation period of about 30 days and a brief pelagic dispersal phase.

conservation status

Not listed by the IUCN. This species was caught for the local market up to the 1950s, when stocks were reported to be over-fished. There is currently no commercial fishery for this species, nor has it been assigned a quota under New Zealand's quota management system.

significance to humans

This is said to be a palatable species, but its fearsome appearance and coarse texture relegate its use mainly to bait in lobster pots.

Emerald notothen

Trematomus bernacchii

family

Nototheniidae

taxonomy

Trematomus bernacchii Boulenger, 1902, Cape Adare and Duke of York Island, North Victoria Land, Antarctica.

other common names

English: Emerald rockcod, bernak; French: Bocasson émeraude; Russian: Trematom-pestryak; Spanish: Austrobacalao esmerelda.

physical characteristics

Reaches a maximum length of 12 in (30 cm) and a maximum weight of 12.3 oz (350 g). It is thick-bodied (length/depth ratio, about 3:9) and light brown to pinkish brown on the sides, with irregular darker blotches; the belly is silvery gray. It is covered with ctenoid scales, except for a patch between the eyes, where there may be a single row of scales. Three distinct color varieties are known:

• continuous white band across the nape and opercula
• an interrupted band, consisting of a narrow medial stripe on the nape and on each operculum
• no white marking

distribution

Coastal waters of Antarctica, including the Antarctic Peninsula and South Shetland and South Orkney Islands.

habitat

It is most common on boulder, rock, or gravel bottoms or in sponge beds at depths of 18–1,200 ft (5–370 m), but it has been caught as deep as 2,280 ft (700 m). Smaller individuals are found in shallow water, whereas mature fish are more common in deeper water.

behavior

The emerald notothen is a sedentary demersal species, spending most of its time immobile on the bottom. It swims infrequently and slowly, sculling with the pectoral fins and making short excursions across the bottom or ascending about 3 ft (1m) into the water column. In the aquarium, dominant individuals establish and defend territories.

feeding ecology and diet

This is an opportunistic feeder and scavenger, primarily on benthic invertebrates. Small crustaceans are the most common dietary item, followed by polychaetes, molluscs, fish, and fish eggs. It ingests seaweed, but its nutritional contribution is unknown.

reproductive biology

The life span of the emerald notothen is 8–10 years; in McMurdo Sound, first spawning occurs at five years. Females spawn biennially; initial oocyte development takes one year and active vitellogenesis (deposition of yolk within the growing egg) another year. Spawning takes place over a brief period, from October to November in Adélie Land and December to January in McMurdo Sound. Fecundity is on the order of 1,500– 3,000 eggs, which are attached to rocks, algae, or sponges; mature eggs are 0.16 in (4 mm) in diameter. There are reports that the emerald notothen tends egg masses in sponges.

conservation status

Not threatened. The emerald notothen is widespread and common; considering its small size and high latitude, it is not likely to come under any significant pressure from fisheries. Its limited mobility probably would lead to extremely rapid depletion of any local populations that might be subjected to commercial fishing.

significance to humans

This species has played an important scientific role in the study of anatomical, physiological, and biochemical adaptations to the Antarctic environment. Most significantly, it has featured prominently in the elucidation of biological antifreeze compounds.

Resources

Books

di Prisco, G., B. Maresca, and B. Tota, eds. Biology of Antarctic Fish. Berlin: Springer-Verlag, 1991.

di Prisco, G., E. Pisano, and A. Clarke, eds. Fishes of Antarctica: A Biological Overview. Milan: Springer-Verlag, 1998.

Eastman, J. T. Antarctic Fish Biology: Evolution in a Unique Environment. San Diego: Academic Press, 1993.

Gon, O, and P. C. Heemstra, eds. Fishes of the Southern Ocean. Grahamstown, South Africa: J. L. B. Smith Institute of Ichthyology, 1990.

Kock, Karl-Hermann. Antarctic Fish and Fisheries. Cambridge: Cambridge University Press, 1992.

Miller, Richard Gordon. History and Atlas of the Fishes of the Antarctic Ocean. Carson City, NV: Foresta Institute for Ocean and Mountain Studies, 1993.

Pisano, E., C. Ozouf-Costaz, C. Bonillo, and F. Mazzei "Pathways of Chromosomal Change During Evolution of Notothenioid Fishes." In Antarctic Ecosystems: Models for Wider Ecological Understanding, edited by W. Davison, C. Howard-Williams, and P. Broady. Christchurch: Caxton Press, 2000.

Periodicals

Chen, L., A. L. DeVries, and C. H. C. Cheng. "Evolution of Antifreeze Glycoprotein Gene from a Trypsinogen Gene in Antarctic Notothenioid Fish." Proceedings of the National Academy of the USA 94 (April 1997): 3811–3816.

Eastman, J. T. "Antarctic Notothenioid Fishes as Subjects for Research in Evolutionary Biology." Antarctic Science 12(2000): 276–287.

Eastman, J. T., and R. R. Eakin. "An Updated Species List for Notothenioid Fish (Perciformes: Notothenioidei), with Comments on Antarctic Species." Archive of Fishery and Marine Research 48 (2000): 11–20.

Eastman, J. T., and A. R. McCune. "Fishes on the Antarctic Continental Shelf: Evolution of a Marine Species Flock?" Journal of Fish Biology 57, suppl. A (2000): 84–102.

Kock, K.-H., and I. Everson. "Biology and Ecology of Mackerel Icefish, Champsocephalus gunnari: An Antarctic Fish Lacking Hemoglobin." Comparative Biochemistry and Physiology 118A, no. 4 (1997): 1067–1077.

Lack, M. "Antarctic Toothfish: An Analysis of Management, Catch and Trade." TRAFFIC Oceania (2001): 1–27.

Lack, M., and G. Sant. "Patagonian Toothfish: Are Conservation and Trade Measures Working?" TRAFFIC Bulletin 19, no. 1 (2001): 1–18.

Montgomery, J., and K. Clements. "Disaptation and Recovery in the Evolution of Antarctic Fishes." Trends in Ecology and Evolution 15 (2000): 267–271.

Stankovic, A., K. Spalik, E. Kamler, et al. "Recent Origin of Sub-Antarctic Notothenioids." Polar Biology 25 (2002): 203–205.

Zhao, Y., M. Ratnayake-Lecamwasam, S. K. Parker, et al. "The Major Adult α-Globin Gene of Antarctic Teleosts and Its Remnants in the Hemoglobinless Icefishes: Calibration of the Mutational Clock for Nuclear Genes." Journal of Biological Chemistry 273, no. 24 (1998): 14745–14752.

Organizations

Australian Society for Fish Biology. 123 Brown Street (PO Box 137), Heidelberg, Victoria 3084 Australia. Phone: 61 (3) 9450 8669. Fax: 61 (3) 9450 8730. E-mail: john.koehn@nre.vic.gov.au Web site: <http://www.asfb.org.au>

Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR). PO Box 213, North Hobart, Tasmania 7002 Australia. Phone: 61 (3) 6231 0366. Fax: 61(3) 6234 9965. E-mail: ccamlr@ccamlr.org Web site: <http:/www.ccamlr.org>

TRAFFIC International. 219c Huntington Road, Cambridge, CB3 0DL United Kingdom. Phone: 44 (0) 1223 277427. Fax: 44 (0) 1223 277237. E-mail: traffic@trafficint.org Web site: <http://www.traffic.org>

Other

Prof J. T. Eastman, Ohio University. "Antarctic Fishes." (March 27, 2003) <http://www.oucom.ohiou.edu/dbms-eastman/index.htm>

John A. Macdonald, PhD