Antarctic icefishes (Channichthyidae): a unique family of fishes. A review, Part II

Icefish or white- blooded fish are a family of species unique among vertebrates in that they possess no haemoglobin. With the exception of one species which occurs on the southern Patagonian shelf, icefish live only in the cold-stable and oxygen-rich environment of the Southern Ocean. It is still questionable how old icefish are in evolutionary terms: they may not be older than 6 Ma, i.e. they evolved well after the Southern Ocean started to cool down or they are 15–20 Ma old and started to evolve some time after the formation of the Antarctic Circumpolar Current. Individuals of most icefish species with the exception of species of the genus Champsocephalus have been found down to 700–800 m depth, a few even down to more than 1,500 m. Icefish have been shown to present organ-level adaptations on different levels to compensate for the ‘disadvantages’ of lacking respiratory pigments. These include a low metabolic rate, well perfused gills, increased blood volume, increased cardiac output, cutaneous uptake of oxygen, increased blood flow with low viscosity, enlarged capillaries, large heart, and increased skin vascularity. Biological features, such as reproduction and growth, are not unique and are comparable to other notothenioids living in the same environment. Icefish produce large yolky eggs which have a diameter of more than 4 mm in most species. Consequently, the number of eggs produced is comparatively small and exceeds 10,000–20,000 eggs in only a few cases. With the exception of species of the genus Champsocephalus which mature at an age of 3 to 4 years, icefish do not attain maturity before they are 5–8 years old. Spawning period of most icefish species is autumn–winter. The incubation period spans from 2 to 3 months in the north of the Southern Ocean to more than 6 months close to the continent. Growth in icefish to the extent it is known is fairly rapid. They grow 6–10 cm in length per annum before they reach spawning maturity. Icefish feed primarily on krill and fish. Some icefish species were abundant enough to be exploited by commercial fisheries, primarily in the 1970s and 1980s with Champsocephalus gunnari as the main target species. Most stocks of this species had been overexploited by the beginning of the 1990s, some had further declined due to natural causes. Other species taken as by-catch species in fisheries were Chaenocephalus aceratus, Pseudochaenichthys georgianus, and Chionodraco rastrospinosus. Chaenodraco wilsoni was the only species exploited on a commercial scale in the high-Antarctic. Part I was published in the preceding issue of Polar Biology. DOI 10.1007/s00300-005-0019-z.     

Two species of antarctic icefishes (genus <Emphasis Type="Italic">Champsocephalus</Emphasis>) share a common genetic lesion leading to the loss of myoglobin expression

Species of the suborder Notothenioidei dominate the fish fauna of coastal Antarctic waters. Members of one notothenioid family, Channichthyidae (Antarctic icefishes), are unique among all vertebrates in lacking the circulating oxygen-binding protein hemoglobin. Icefish species also do not uniformly express the intracellular oxygen-binding protein myoglobin (Mb) in their oxidative muscles. Our laboratory previously characterized the pattern of cardiac Mb expression in 13 of the 16 known icefish species. In this paper, we complete the survey of cardiac Mb expression among all 16 known species of icefishes. Using PAGE and immunoblot analyses, we demonstrate that both Channichthys rhinoceratus and Cryodraco atkinsoni express Mb in heart ventricle, while Champsocephalus esox does not express the protein. We report Mb gene sequences from Channichthys rhinoceratus and Champsocephalus esox genomic DNA. The Mb gene of C. esox contains the identical 5-bp duplication/insertion to that observed in congeneric Champsocephalus gunnari, a species that also does not produce Mb. This duplication in exon 2 of the Champsocephalus spp. gene causes a shift in reading frame at a position normally encoding for amino acid 91 and also results in a premature stop codon, thus disrupting translation of the normal protein. Thus, 6 of the 16 known icefish species do not express cardiac Mb. These results confirm earlier conclusions that losses of Mb expression have occurred via at least four independent events during the evolution of the icefish family. Extreme similarity of Mb genes in Champsocephalus congeners further suggests recent speciation despite early divergence of this group from the lineage leading to more derived icefishes.     

Drinking in Antarctic fishes

Drinking rates have never been measured in Antarctic fish. Drinking rates were measured in four species of notothenioid fish, including a hemoglobinless icefish, found in the near-freezing waters of the Ross Sea of the Southern Ocean. All of the fish, with the exception of the icefish, had low drinking rates and high serum osmolalities relative to temperate seawater fish. The icefish had significantly higher drinking rates and serum osmolalities relative to the Antarctic fishes containing hemoglobin, including Trematomus bernacchii. Warm acclimation of T. bernacchii, from −1.5°C to +4°C for 4 weeks, significantly increased their drinking rates 4.6-fold, significantly decreased their serum and intestinal osmolality by 11% and 12%, respectively, relative to cold-acclimated fish. These results indicate that increased drinking rates in Antarctic fish at elevated temperatures are involved in maintaining a lower serum osmolality.     

Degradation of icefishes (Salangidae) in the Yangtze River basin of China: threats and strategies

The Yangtze River basin has been a major area in terms of icefish diversity conservation and fisheries. In the past two to five decades, however, the wild icefish resources have degraded continuously in both the brackish estuary and freshwater portions of the basin. Stocks have declined in all icefish fisheries, resulting in an annual fishing yield decrease from the maximum of 4,000 metric tons to less than 1,300 metric tons in this river basin. The stocks decline significantly (t = −6.74, df = 1, p < 0.001) correlated with icefish fishing stress in the estuary. Meanwhile, species composition changed markedly with smaller sized icefish species increased relative to the number of larger ones in terms of their relative abundances. Following the stocks decline and size reduction, icefish commercial value decreased to the point that 91.9% of the traditional icefish-producing waters (57 out of 62) terminated their icefish fisheries and some species became locally extinct. Based on the degradation process, we suggest that these changes in stock size and species diversity are the result of overfishing and habitat loss and degradation. We propose a number of measures aimed at the conservation and restoration of the wild icefish resources in the Yangtze River basin.     

Fisheries in the Southern Ocean: an ecosystem approach

The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) is bound by its Article II, 3 to follow an ecosystem approach to management. This approach has been extended to the application of a precautionary approach in the late 1980s. In our review, we deal primarily with the science-related aspects of CCAMLR and its development towards an ecosystem approach to the management of the living resources of the Southern Ocean. To assist the Commission in meeting its objectives, as set out in Article II, 3, the Scientific Committee established the CCAMLR Ecosystem Monitoring Programme to detect possible effects of krill fishing on the performance of top-level predators, such as albatrosses, penguins, petrels and fur seals. Fisheries in the Southern Ocean followed the fate of other fisheries worldwide in which target species were depleted to low level one after the other. Currently, two types of fisheries are open: the longline fisheries on Patagonian toothfish (Dissostichus eleginoides) and Antarctic toothfish (Dissostichus mawsoni) and the trawl fisheries on mackerel icefish (Champsocephalus gunnari). Both fisheries are managed in a single-species context, however, with conservation measures in place to protect by-catch species, such as rattails (Macrouridae) and skates and rays (Rajidae). Two major problems still exist in fisheries in the Southern Ocean: the by-catch of birds in longline fisheries primarily in the Indian Ocean and the high level of IUU fishing again in the Indian Ocean. Both, the by-catch of birds and high IUU catches undermine the credibility of CCAMLR to safeguard the marine living resources in the Southern Ocean.     

Competition, mortality and diversity in South Atlantic coastal boulder communities

Boulder shores are common at all latitudes and dominate the intertidal and subtidal zones of sub-Antarctic coastlines. The encrusting benthos of boulders was examined on similar shore types at four locations: Tierra del Fuego, East Falkland, West Falkland and Bird Island (off South Georgia). Bird Island is unusual in experiencing high trampling and organic enrichment from fur seals. The results were compared to a Patagonian site and a non-trampled South Georgia site (Husvik) and other sites taken from the literature. Principal Component Analysis revealed South Atlantic/Southern Ocean encrusting faunas formed a distinct cluster when compared to assemblages from elsewhere at similar latitudes. Bray Curtis cluster analysis of the South Atlantic-Southern Ocean sites showed the major division was between Southern Ocean and South Atlantic Ocean sites, beyond which there were three distinct clusters centred around Patagonia (Magellanic), the Falklands and Southern Ocean sites. The organisation of competitive interactions between species was mostly determinate and transitive (essentially hierarchical). The transitivity index scores were higher than most similar assemblages studied to date. The diversity of encrusting assemblages ranged from Shannon Weaver H values of 2.38–0.77 (East Falkland and Bird Island, respectively) in the intertidal to 1.27–0.73 (Patagonia and South Georgia, respectively) in the subtidal zone. Annual mortality (of bryozoan colonies) varied from 85–97% in the intertidal to 65–92% in the subtidal, being higher in the Southern Ocean than South Atlantic sites, largely due to ice scour and wave action. The Bird Island mortality values may be high even for a Southern Ocean site. Accepted: 5 October 2000     

Determining the frequency of heteropteran predation on sweetpotato whitefly and pink bollworm using multiple ELISAs

The gut contents of field-collected, predaceous Heteroptera were assayed for the presence of eggs of the sweetpotato whitefly,Bemisia tabaci Gennadius (Homoptera: Aleyrodidae) and the pink bollworm,Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae) using multiple enzyme-linked immunosorbent assays (ELISAs). Of seven species examined,Geocoris species andOrius tristicolor (Say) were the most frequent predators of sweetpotato whitefly with 32–39% of the individuals tested over the whole season scoring positive for whitefly antigens. With the exception ofLygus hesperus Knight, a major insect pest as well as a predator, the frequency of predation on pink bollworm eggs was much lower (0.7–14.3% positive over the season). Relatively few predators tested positive for both antigens (0.3–12.5%).     

Growth and reproduction of the non‐native icefish Neosalanx taihuensis Chen, 1956 (Salangidae) in a plateau lake,southwestern China

Growth, reproduction and abundance traits of the invasive icefish Neosalanx taihuensis Chen, 1956 were investigated monthly from July 2009 to May 2011 in Lake Erhai on the Yunnan‐Guizhou Plateau, south‐western China, in order to explore the changes in life‐history traits after translocation. The results indicated that the icefish exhibited obvious plasticity in growth and reproduction traits. Growth of the fish in Lake Erhai was faster than that in native waters and in other translocated reservoirs. By fitting the von Bertalanffy growth model to the data, it was estimated that icefish obtain an asymptotic size of 96.12 mm, a K of 1.61, and a t₀ of ‐0.26; the calculated overall growth performance index φ′ was 4.17. The strategy of reproduction changed from multiple‐ to single‐spawning. The spawning period was from October to December with the absolute and relative fecundities of 1250 ± 169 eggs per ind and 2557 ± 245 eggs per g, respectively. Plasticity in icefish growth and reproduction in Lake Erhai greatly facilitated its population establishment, making it one of the most abundant fish species. The icefish invasion in the lake may be one of the reasons for the decrease or extinction of native fish species populations, and some measures for the control of this invasive fish are suggested.     

Neoascarophis macrouri n. sp. (Nematoda: Cystidicolidae) from the stomach of Macrourus berglax (Macrouridae) in the eastern Greenland Sea

A new species of parasitic nematode, Neoascarophis macrouri n. sp. (Cystidicolidae), is described from the stomach and stomach wall of the marine deep-water fish Macrourus berglax (onion-eye grenadier) in the eastern Greenland Sea (North Atlantic Ocean). The new species, studied using both light and scanning electron microscopy, is characterised mainly by the location of the vulva near the posterior end of the body (a short distance anterior to the anus), non-filamented eggs, the structure of the mouth, a short vestibule and the length of the spicules (567–615 and 144–156 μm). Metabronema insulanum Solov’eva, 1991 is transferred to Neoascarophis as N. insulana (Solov’eva, 1991) n. comb.     

Temporal and spatial variation of dominant pelagic Copepoda (Crustacea) in the Weddell Sea (Southern Ocean) 1929 to 1993

The abundances of four dominant Antarctic copepod species, Metridia gerlachei, Rhincalanus gigas, Calanoides acutus and Calanus propinquus, were examined in the Southern Ocean in a combination of a literature review, analysis of museum samples and field sampling. The data were analysed for spatial and temporal variations. The data included in the analysis were from the Weddell Sea area in the summertime at periods 1929–1939 and 1989–1993. The results are discussed in the light of environmental changes and their hypothesised and observed consequences in the Southern Ocean: global temperature change, ozone deficiency and cascading trophic interactions. Combining all these hypothetical effects our null hypothesis was that there were no consistent long-term changes in the abundance of dominant pelagic Copepoda. The null hypothesis was rejected, since several taxons did show statistically significant long-term changes in abundance. The changes were not uniform however. The numbers of adults and juveniles of Calanus propinquus increased significantly between the periods studied. Adult stages of Calanoides acutus were the only taxon decreasing in abundance, in concert with the cascading trophic interactions theory. Latitudinally, only Metridia gerlachei showed a significant increase from north to south. Longitudinally, the abundances of Calanus propinquus juveniles and both adults and juveniles of Rhincalanus gigas increased from west to east. There were no significant variations between day and night samples. Interannual changes were statistically significant in juvenile stages of all the species and in adults of Calanus propinquus. We conclude that no uniform and consistent abundance changes could be observed in the pelagic Copepoda of the Weddell Sea that could be connected to major environmental changes, expected to affect the whole planktonic ecosystem of the Southern Ocean. Significant changes in some of the species studied show that the pelagic ecosystem is not in a steady state, but in addition to interannual changes, there are also major fluctuations extending over decades. Received: 5 December 1996 / Accepted: 24 March 1997     

Construction, electroporatic transfer and expression of ZpβypGH and ZpβrtGH in zebrafish

Recombinant transformation vectors (ZPβypGH and ZpβrtGH) consisting of fish growth hormone cDNA, and a reporter geneβ-galactosidase driven by fish promoter (Zp) were constructed. Freshly fertilized eggs of zebrafish (Brachydanio rerio) were electroporated at optimum conditions (0.07 kV voltage; 25 μF capacitance; 8 ohm resistance and 2 pulses) in the presence of one of these transformation vectors (100 μg circular DNNml). In either cases 72% of the electroporated eggs successfully hatched, in comparison to the 85% hatchability of the control eggs. Genomic DNA extracted from fins of randomly chosenF ₀ individuals was screened (by Southern blot hybridization); the transgenes were retained in the host genome of all the randomly chosen adult transformants. Fin-positive presumptive founder parents were crossed with control counterparts and the DNA of randomly chosenF ₁ progenies was screened for germline transformation. Southern analysis of chosenF ₁ progenies revealed the persistence of ZPβypGH or ZpβrtGH in 53% of theF ₁ progenies. Southern analyses of chosenF ₁ progenies and the frequency (53% ofF ₁ ZpβrtGH and 53% ofF ₁ ZP{β}ypGH) of transmission revealed the degree of mosaicism inF ₀ transformants. Expression was confirmed from the 3–4 times elevated levels of activity of the reporter gene and 30–40% accelerated growth of transgenicF ₀ andF ₁ progenies. Construction of the transgenes was made at the Taiwan National Ocean University, Taiwan and all other work at the Madurai Kamaraj University, Madurai.     

Comparison of the ovaries and oogenesis of some Australian and South African viviparid clinid fishes (Clinidae,Blennioidei, Perciformes)

The ovaries of the studied 16 viviparous blenny species of the fish family Clinidae (Teleostei, Perciformes) from Australia and South Africa, are of the cystovarian type and bilobed, ending in a common short oviduct at the gonopore; they are elongated in juvenile fish, becoming bottle-shaped or oval with maturation and onset of embryogenesis, when they comprise 18–20% of the total body weight. The ovarian lumen is covered by the germinal epithelium, resting on a basal membrane. The primordial germ cells are situated along the inner side of each lobe, juxtaposed internally to the lumenal epithelium. Single primary oocytes or nests of such, each in a follicular envelope and covered externally by the lumenal epithelium forming clusters protruding into the lumen, along which the inoculated sperm settle. Two different types of reproductive strategies are demonstrated by the Australian species of Heteroclinus and Cristiceps and the South African species of Clinus and related forms. The Australian species produce high numbers of small (350 ± 50 μm in diameter) eggs that develop synchronously, while the South African species produce much fewer and larger (1.0 ± 0.1 mm) eggs, asynchronous in development. An exception in the latter group is Blennophis striatus, with eggs of ca.1.4 mm that develop synchronously. The Australian species are thus mass releasers that bear and release up to 1000 small (11 ± 1.5 mm in length) embryos/spawn, while the South African Clinus species are discrete releasers that bear eggs of various sizes and small numbers of mature for release embryos that reach up to 22 mm in length, and are released throughout the year. The cytogenesis of the oocytes and surrounding envelopes is discussed.     

Dolichopteryx rostrata, a new species of spookfish (Argentinoidea: Opisthoproctidae) from the eastern North Atlantic Ocean

A new species, Dolichopteryx rostrata, is described on the basis of a single specimen (66.2 mm in standard length) collected west of the Hebrides Islands, eastern North Atlantic Ocean. The new species is characterized by an elongate snout and head, small pouchlike eyes, an adipose fin, short dorsal fin base, anal fin base originating under dorsal fin base, a clear longitudinal suborbital brownish band extending forward from behind posterior margin of orbit to snout tip, and 41 (=26 + 15) vertebrae. Total fecundity is low; the ovarian eggs number only 473, despite the ovary having developed ova. Ovarian eggs could be clearly subdivided into an “undeveloped group” (0.1–0.7 mm diameter classes, n = 405) and a “developed group” (0.9–1.3 mm classes, n = 68), based on their frequency distribution. Supplementary material to this paper is available in electronic format at http://dx.doi.org/10.1007/s10228-005-0306-2     

Sexual reproduction in three hermaphroditic deep-sea Caryophyllia species (Anthozoa: Scleractinia) from the NE Atlantic Ocean

The reproductive biology and gametogenesis of three species of Caryophyllia were examined using histological techniques. Caryophyllia ambrosia, Alcock 1898, C. cornuformis, Pourtales 1868, and C. sequenzae, Duncan 1873, were collected from the Porcupine Seabight and Rockall Trough in the NE Atlantic Ocean. These three ahermatypic solitary corals inhabit different depth ranges: C. cornuformis – 435–2000 m, C. sequenzae – 960–1900 m, and C. ambrosia – 1100–3000 m. All three species are hermaphroditic. Hermaphroditism in these species was found to be cyclical, with only one sex of gametes viable in any individual at any point in time, although gametes of both sexes were found together within a single mesentery. Once the viable gametes are spawned, the next sex of gametes continues to grow until mature, and so gametogenesis is a continuous cycle. Oocytes and spermacysts in all species increased in density towards the actinopharynx. Maximum fecundity for C. sequenzae was 940 oocytes per polyp, and for C. ambrosia 2900 oocytes per polyp. Fecundity could not be established for C. cornuformis. In all three species, individuals were asynchronous within populations, and production of gametes was quasi-continuous throughout the year. All species are hypothesised to have lecithotrophic larvae owing to their large oocyte sizes (C. cornuformis max – 350 μm; C. sequenzae max – 430 μm; C. ambrosia max – 700 μm). Both the average oocyte size and fecundity increased in species going down the depth gradient of the NE Atlantic.     

Response ofChelonus blackburni [Hym.: Braconidae] to different ages and densities of potato tuber moth eggs

The response of an efficient egg-larval parasitoidChelonus blackburni Cameron to different ages and densities of potato tuber mothPhthorimaea operculella (Zeller) eggs was studied in a laboratory system. The parasitoid accepted 0–1, 1–2 and 2–3 day old eggs for parasitization. However, the eggs belonging to the first two age groups were significantly more acceptable when compared to the last age group. At low host density levels of 2 and 5 eggs per parasitoid, parasitoid emergence was zero. On increasing the host density, percent progeny production increased till a plateau was reached at a parasitoid-host ratio of 1∶50. This ratio thus proves to be ideal for obtaining maximum percent progeny production in the case of 0–1 and 1–2 day old eggs. Further increase in host density did not result in any significant increase in percent progeny production.      

The biology of the spiny icefish <Emphasis Type="Italic">Chaenodraco wilsoni</Emphasis> Regan, 1914

The most abundant ice fish species observed in catches off the northern tip of the Antarctic Peninsula in the last 25–30 years has been the spiny ice fish Chaenodraco wilsoni Regan 1914. C. wilsoni has been exploited on a commercial scale from the late 1970s to the end of the 1980s off Joinville–D’Urville Islands (CCAMLR Statistical Subarea 48.1) and in the Cosmonauts and Cooperation Seas and Prydz Bay in the Indian Ocean sector (CCAMLR Statistical Division 58.4.2). This paper presents new information on biological features and life history characteristics of C. wilsoni, based on research survey collections along the northern Antarctic Peninsula in 2006 and 2007 and samples taken in the commercial fishery in 1987. Length frequency compositions from the research surveys demonstrated that fish 21–34 cm long predominated in the catches. Sexual maturity is attained at 24–25 cm. Absolute fecundity and relative fecundity is low (1,000–2,500 eggs; 6–12 eggs). Oocyte diameter varied from 4.0 to 4.9 mm very close to spawning. Spawning at the tip of the Antarctic Peninsula is likely to occur in October–November. Remotely operated vehicle deployments in the northern Weddell Sea demonstrated that C. wilsoni exhibit parental nest guarding where males protect the eggs. The incubation period is likely to be 8 months long. Fish feed primarily on Antarctic krill (Euphausia superba) in the Antarctic Peninsula region and in the Cosmonauts and Cooperation Seas while fish take ice krill (Euphausia crystallorophias), Pleuragramma antarcticum and myctophids to some extent in other areas. Age determination still awaits validation. Preliminary ageing attempts suggested a maximum age of about 8–10 years.     

The influence of host's age on the selectivity of parasitism and fecundity ofTrichogramma

Laboratory studies showed that ♀♀ ofTrichogramma embryophagum (Hartig) andT. semblidis (Aurivillius) when offered an alternative of young and old eggs of the grain moth,Sitotroga cerealella Oliv., prefer ovipositing into young eggs. The fecundity of ♀♀ that are simultaneously offered both young and old eggs was lower in comparison to ♀♀ offered only young eggs (inspite of the fact that in both variants young eggs were in excess). The results suggest that old eggs can inhibit the oviposition of some species ofTrichogramma.      

Developmental morphology of the Indian cyprinid fish Barilius canarensis

Embryonic and larval development of an Indian cyprinid fish, Barilius canarensis, is described from laboratory-reared specimens. The eggs, measuring 2.1–2.4 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yellow yolk without oil globules. Hatching occurred 39–45 h after fertilization at 26.8°–27.4°C. The newly hatched larvae, measuring 4.8–5.1 mm in body length (BL) with 22 + 17 = 39 myomeres, were characterized by melanophores already deposited on the eyes. The eggs of B. canarensis resembled those of the related danionin species Candidia barbatus, Opsariichthys uncirostris uncirostris, Zacco platypus, Z. sieboldii, and Z. temminckii. Although the larvae of B. canarensis were also similar to those of the foregoing species in general morphology, they differed in having a straight notochord tip and pigmentation on the eyes at hatching and the almost entire absence of melanophores on the ventral body surface from the yolk sac to postflexion larval stages. Conversely, melanophores occurred on the anterior abdominal and pericardial cavities from the preflexion to postflexion larval stages.     

Systematics and distribution of theDiplophos taenia species complex (Gonostomatidae), with a description of a new species

For the systematic study of the gonostomatid fishes of theDiplophos taenia species complex, a total of 698 specimens was obtained from the three oceans. Four valid species were recognized:D. taenia Günther,D. proximus Parr,D. orientalis Matsubara, andD. australis sp. nov. The diagnostic characters are 90–100 total vertebrae (TV) (37–41 abdominal (AV) +52–60 caudal vertebrae (CV)) and 103–115 IC photophores (IC) for D.taenia, 85–90 TV (36–39 AV + 48–52 CV) and 98–104 IC forD. proximus, 83–86 TV (33–35 AV + 49–52 CV) and 92–100 IC for D.orientalis, and 84–91 TV (33–37 AV+ 50–54 CV) and 99–105 IC forD. australis. In addition to the above,D. proximus has larger orbital diameter (the proportion to head length 21–28%) than the other three species (15–23%) beyond 70mm in standard length. Due to wide distribution,D. taenia shows meristic variations: the numbers of TV, IC and anal fin rays (A) are smaller at lower latitudes and larger at higher ones in all oceans, and the number of A is smaller in the Atlantic (56–71) than in the other oceans (59–72) of the same latitude. Because of these variations, identification to species level is possible only area by area. The distribution of each of the four species is also distinct:D. taenia is a cosmopolitan between about 40° N and 30° S exclusive of the eastern tropical Pacific;D. proximus is endemic to the eastern tropical Pacific;D. orientalis is limited to the North Pacific transitional zone between about 30° and 40° N; andD. australis in a transitional zone of the Southern Ocean south of 20° S.