A description of the first intact Dwarf Sperm whale from the South China Sea and a review of documented specimens of the Kogiidae (Cetacea) from Hong Kong

Reports of the Dwarf Sperm whale (Kogia sima) in Chinese waters exist only for four specimens, which stranded along the coast of the East China Sea. Records of this species in the South China Sea consist of one skull noted as present in a Vietnamese ‘whale temple’. Recently recognized material in the biological sciences building of the University of Hong Kong, confirms the presence of both the Dwarf (K. sima) and the Pygmy (K. breviceps) Sperm whales in Hong Kong. In addition, close examination of Hong Kong stranding records reveals that of the four specimens identified as K. breviceps one is, in fact, K. sima. A comparison of the Hong Kong material, the holotype of K. sima and other Asian specimens of Kogia spp. affirms that most of the features already used elsewhere to distinguish the two species are reliable.     

Domoic acid exposure in pygmy and dwarf sperm whales (Kogia spp.) from southeastern and mid-Atlantic U.S. waters

The neurotoxin domoic acid (DA) was detected in urine and fecal samples recovered from pygmy sperm whales (Kogia breviceps) and dwarf sperm whales (Kogia sima) stranding along the U.S. Atlantic coast from 1997 to 2008. Of the 41 animals analyzed from Virginia, North Carolina, South Carolina and Florida, 24 (59%) tested positive for DA at concentrations of 0.4–1.8 ng/mL in urine and 12–13,566 ng/g in feces as determined by liquid chromatography–tandem mass spectrometry (LC–MS/MS). Feces appeared to be the best indicator of DA exposure in Kogia spp., with 87% of all fecal samples analyzed testing positive for this toxin. Additional stranded animals (n = 40) representing 11 other cetacean species were recovered from the same region between 2006 and 2008 and analyzed by LC–MS/MS, however DA was not detected in any of these individuals. DA is produced naturally by diatoms in the genus Pseudo-nitzschia. Although blooms of DA-producing Pseudo-nitzschia have been associated with repeated large-scale marine mammal mortalities on the west coast of the U.S., there is no documented history of similar blooms on the southeast U.S. coast, and there were no observed Pseudo-nitzschia blooms in the region associated with any of these strandings. The feeding habits of Kogia spp. are poorly documented; thus, the vector(s) for DA exposure to these deep-diving species remains to be identified. Toxin accumulation in these pelagic whale species may be an indication of cryptic harmful algal bloom activity in offshore areas not currently being monitored. This study highlights the need for a better understanding of the role of toxigenic algae in marine mammal morbidity and mortality globally.     

Evolutionary Patterns among Living and Fossil Kogiid Sperm Whales: Evidence from the Neogene of Central America

Kogiids are known by two living species, the pygmy and dwarf sperm whale (Kogia breviceps and K. sima). Both are relatively rare, and as their names suggest, they are closely related to the sperm whale, all being characterized by the presence of a spermaceti organ. However, this organ is much reduced in kogiids and may have become functionally different. Here we describe a fossil kogiid from the late Miocene of Panama and we explore the evolutionary history of the group with special attention to this evolutionary reduction. The fossil consists of cranial material from the late Tortonian (~7.5 Ma) Piña facies of the Chagres Formation in Panama. Detailed comparison with other fossil and extant kogiids and the results of a phylogenetic analysis place the Panamanian kogiid, herein named Nanokogia isthmia gen. et sp. nov., as a taxon most closely related to Praekogia cedrosensis from the Messinian (~6 Ma) of Baja California and to Kogia spp. Furthermore our results show that reduction of the spermaceti organ has occurred iteratively in kogiids, once in Thalassocetus antwerpiensis in the early-middle Miocene, and more recently in Kogia spp. Additionally, we estimate the divergence between extant species of Kogia at around the late Pliocene, later than previously predicted by molecular estimates. Finally, comparison of Nanokogia with the coeval Scaphokogia cochlearis from Peru shows that these two species display a greater morphological disparity between them than that observed between the extant members of the group. We hypothesize that this reflects differences in feeding ecologies of the two species, with Nanokogia being more similar to extant Kogia. Nanokogia shows that kogiids have been part of the Neotropical marine mammal communities at least since the late Miocene, and gives us insight into the evolutionary history and origins of one of the rarest groups of living whales.     

The diet of pygmy sperm whales, Kogia breviceps, stranded in New Zealand: implications for conservation

The stomach contents of 27 pygmy sperm whales, Kogia breviceps, stranded on New Zealand beaches between 1991 and 2003 are reported. These individuals comprise 16 males, 10 females, and one for which no sex information is available. The diet was found to include fish and crustaceans, but is comprised primarily of cephalopods, with 0–526 lower beaks, representing an estimated maximum of c. 60 kg of cephalopod prey consumed by any one whale. Cephalopod prey is attributed to 23 species from 13 families, and is dominated by juvenile individuals of the families Histioteuthididae and Cranchiidae (adults of which usually occur at depths exceeding 400 m). Perceived threats to this whale, particularly those affecting distribution and abundance of prey species, are also discussed. These are the first data reporting the diet of this whale species in New Zealand waters. A comparison of the diet of K. breviceps is made with that of the sperm whale, Physeter macrocephalus from New Zealand waters, and with the diet of Kogia known elsewhere.     

Detection of the elusive Dwarf sperm whale (Kogia sima) using environmental DNA at Malpelo island (Eastern Pacific,Colombia)

1. Monitoring large marine mammals is challenging due to their low abundances in general, an ability to move over large distances and wide geographical range sizes.
2. The distribution of the pygmy (Kogia breviceps) and dwarf (Kogia sima) sperm whales is informed by relatively rare sightings, which does not permit accurate estimates of their distribution ranges. Hence, their conservation status has long remained Data Deficient (DD) in the Red list of the International Union for Conservation of Nature (IUCN), which prevent appropriate conservation measures.
3. Environmental DNA (eDNA) metabarcoding uses DNA traces left by organisms in their environments to detect the presence of targeted taxon, and is here proved to be useful to increase our knowledge on the distribution of rare but emblematic megafauna.
4. Retrieving eDNA from filtered surface water provides the first detection of the Dwarf sperm whale (Kogia sima) around the remote Malpelo island (Colombia).
5. Environmental DNA collected during oceanic missions can generate better knowledge on rare but emblematic animals even in regions that are generally well sampled for other taxa.

     

The use of DNA barcoding to monitor the marine mammal biodiversity along the French Atlantic coast

In the last ten years, 14 species of cetaceans and five species of pinnipeds stranded along the Atlantic coast of Brittany in the North West of France. All species included, an average of 150 animals strand each year in this area. Based on reports from the stranding network operating along this coast, the most common stranding events comprise six cetacean species (Delphinus delphis, Tursiops truncatus, Stenella coeruleoalba, Globicephala melas, Grampus griseus, Phocoena phocoena)and one pinniped species (Halichoerus grypus). Rare stranding events include deep-diving or exotic species, such as arctic seals. In this study, our aim was to determine the potential contribution of DNA barcoding to the monitoring of marine mammal biodiversity as performed by the stranding network.We sequenced more than 500 bp of the 5’ end of the mitochondrial COI gene of 89 animals of 15 different species (12 cetaceans, and three pinnipeds). Except for members of the Delphininae, all species were unambiguously discriminated on the basis of their COI sequences. We then applied DNA barcoding to identify some “undetermined” samples. With again the exception of the Delphininae, this was successful using the BOLD identification engine. For samples of the Delphininae, we sequenced a portion of the mitochondrial control region (MCR), and using a non-metric multidimentional scaling plot and posterior probability calculations we were able to determine putatively each species. We then showed, in the case of the harbour porpoise, that COI polymorphisms, although being lower than MCR ones, could also be used to assess intraspecific variability. All these results show that the use of DNA barcoding in conjunction with a stranding network could clearly increase the accuracy of the monitoring of marine mammal biodiversity.     

CETACEAN ABUNDANCE IN HAWAIIAN WATERS ESTIMATED FROM A SUMMER/FALL SURVEY IN 2002

Cetacean abundance is estimated for the U.S. Exclusive Economic Zone (EEZ) around the Hawaiian Islands based on a ship line‐transect survey from August to November, 2002. Sighting detection functions are estimated from this and other NOAA research surveys from 1986 to 2002 using a new, multiple‐covariate approach. Twenty‐four species were seen on this survey, including two species (Fraser's dolphin [Lagenodelphis hosei] and sei whale [Balaenoptera borealis]) that had not been previously documented to occur in Hawaiian waters. The most abundant large whales are sperm whales (Physeter macrocephalus) and Bryde's whales (Balaenoptera edeni). The most abundant delphinids are pilot whales (Globicephala macrorhynchus), rough‐toothed dolphins (Steno bredanensis), Fraser's dolphins, spotted dolphins (Stenella attenuata), and striped dolphins (Stenella coeruleoalba). Dwarf and pygmy sperm whales (Kogia sima and Kogia breviceps) and Cuvier's beaked whales (Ziphius cavirostris) are also estimated to be quite abundant. Some of the migratory baleen whales (fin whales [Balaenoptera physalus], sei whales, minke whales [B. acutorostrata], and humpback whales [Megaptera novaeangliae]) were seen only late in the survey. Abundance is estimated for 19 cetacean species. The overall density of cetaceans is low in the study area, especially for delphinids. The precision of density and abundance estimates is generally low for all species because of the low number of sightings.     

Review of cetaceans stranded on the Irish coast, 1901–95

Published records of cetaceans stranded on the Irish coast during the period 1901–95 are reviewed. In this review the number of stranding events has been used in the analysis and includes both live strandings and those animals washed up dead. There were 529 records involving 21 species. The Harbour Porpoise (27%) was the most frequently reported species, followed by Common Dolphins (16%) and Pilot Whales (15%). Minke Whales (8%) were the most frequently reported mysticete. The number of reported strandings has increased since the 1960s which is thought to be mainly due to increased observer effort. Cetaceans have stranded on all Irish coasts but mainly along the south coast and along the western seaboard but with no apparent overall seasonal trend. There was a peak in the strandings of Common Dolphins during 1991–92 when 27% (28 records) of all strandings were reported and of White-sided Dolphins when 60% (28 records) were reported, both of which were attributed to possible interactions with fisheries. The number of Striped Dolphins stranded on the Irish coast has increased steadily since the 1980s and may reflect increasing water temperatures.
These stranding records are considered inadequate to determine the status of most species of cetaceans in Irish waters but are sufficient to identify unusual stranding events such as high mortalities due to fisheries interactions or epizootics. More observer coverage is required before the stranding data are adequate for monitoring the status of most species but a stranding scheme is considered the most effective and efficient method of long-term monitoring of cetaceans in Irish waters.     

New insights into whale shark <Emphasis Type="Italic">Rhincodon typus</Emphasis> diet in Brazil: an observation of ram filter-feeding on crab larvae and analysis of stomach contents from the first stranding in Bahia state

We describe the first record of a whale shark, Rhincodon typus, feeding in Brazilian coastal waters, and the first stranding record in the state of Bahia, Northeast Brazil. In April 2008, an individual of R. typus was observed surface feeding on Dromiidae crab larvae in the continental shelf off the coast of Bahia, near a gas platform. Other fishes were observed foraging in association with the whale shark. We also document the first stranding of R. typus on the coast of Bahia in October 2013. Biometric data confirmed that the stranded whale shark was a juvenile. Stomach content analysis revealed the ingestion of Geryonidae crab larvae. Plastic debris were also found in the gastric lumen of the stranded juvenile whale shark, and we speculate that it could have been a contributing factor to the stranding, and subsequent death of the whale shark. Crab larvae were observed in both of our records and likely to consist as relevant prey items for R. typus in Brazilian continental shelf. Our study provided a contribution on the diet and feeding behaviour of whale sharks in tropical oligotrophic waters and highlights the risks of marine pollution for the species conservation.     

Prevalence and anatomic site of Crassicauda sp. infection,and its use in species identification,in kogiid whales from the mid‐Atlantic United States

The parasitic nematode Crassicauda sp. was initially described in kogiid whales from specimens collected within cervical tissues, uncommon sites of infection for this parasite. Crassicauda sp. has only been reported in Kogia breviceps to date, but no study has yet investigated a large sample of both kogiid species. A 15 yr record of 104 kogiid strandings (K. sima, n = 40; K. breviceps, n = 64) in North Carolina and Virginia, U.S.A. was used to determine the prevalence of Crassicauda sp. across species, within species across sex, and within sex across length and life history categories. Crassicauda sp. was confirmed to be a species‐specific parasite among kogiids infecting only K. breviceps (prevalence = 45%). Within K. breviceps, prevalence was similar (45%) in both immature and mature males, but increased from 10% in immature to 76% in mature females. This study confirmed the cervico‐thoracic distribution of the parasite, and identified a novel site of infection in a previously undescribed exocrine gland associated with the pigmented “false gill slit.” The species‐specific nature of Crassicauda sp. infection, the exocrine gland, and the distinct features of the false gill slit pigmentation associated with the gland, are all useful characters to identify kogiid species in the field.     

Marine Mammal Strandings and Environmental Changes: A 15-Year Study in the St. Lawrence Ecosystem

Understanding the effects of climatic variability on marine mammals is challenging due to the complexity of ecological interactions. We used general linear models to analyze a 15-year database documenting marine mammal strandings (1994–2008; n = 1,193) and nine environmental parameters known to affect marine mammal survival, from regional (sea ice) to continental scales (North Atlantic Oscillation, NAO). Stranding events were more frequent during summer and fall than other seasons, and have increased since 1994. Poor ice conditions observed during the same period may have affected marine mammals either directly, by modulating the availability of habitat for feeding and breeding activities, or indirectly, through changes in water conditions and marine productivity (krill abundance). For most species (75%, n = 6 species), a low volume of ice was correlated with increasing frequency of stranding events (e.g. R² _adj = 0.59, hooded seal, Cystophora cristata). This likely led to an increase in seal mortality during the breeding period, but also to increase habitat availability for seasonal migratory cetaceans using ice-free areas during winter. We also detected a high frequency of stranding events for mysticete species (minke whale, Balaenoptera acutorostrata) and resident species (beluga, Delphinapterus leucas), correlated with low krill abundance since 1994. Positive NAO indices were positively correlated with high frequencies of stranding events for resident and seasonal migratory cetaceans, as well as rare species (R² _adj = 0.53, 0.81 and 0.34, respectively). This contrasts with seal mass stranding numbers, which were negatively correlated with a positive NAO index. In addition, an unusual multiple species mortality event (n = 114, 62% of total annual mortality) in 2008 was caused by a harmful algal bloom. Our findings provide an empirical baseline in understanding marine mammal survival when faced with climatic variability. This is a promising step in integrating stranding records to monitor the consequences of environmental changes in marine ecosystems over long time scales.     

Foraging ecology and niche overlap in pygmy (Kogia breviceps) and dwarf (Kogia sima) sperm whales from waters of the U.S. mid‐Atlantic coast

A complementary approach of stomach content and stable isotope analyses was used to characterize the foraging ecology and evaluate niche overlap between pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales stranded on the U.S. mid‐Atlantic coast between 1998 and 2011. Food habits analysis demonstrated both species were primarily teuthophagous, with 35 species of cephalopods, and 2 species of mesopelagic fishes represented in their overall diets. Pianka's Index of niche overlap suggested high overlap between whale diets (O_n = 0.92), with squids from the families Histioteuthidae, Cranchidae, and Ommastrephidae serving as primary prey. Pygmy sperm whales consumed slightly larger prey sizes (mean mantle length [ML] = 10.8 cm) than dwarf sperm whales (mean ML = 7.8 cm). Mean prey sizes consumed by pygmy sperm whales increased with growth, but showed no trend in dwarf sperm whales. Significant differences were not detected in δ¹⁵N and δ¹³C values of muscle tissues from pygmy (10.8‰ ± 0.5‰, ?17.1‰ ± 0.6‰), and dwarf sperm whales (10.7‰ ± 0.5‰, ?17.0‰ ± 0.4‰), respectively. Isotopic niche widths also did not differ significantly and dietary overlap was high between the two species. Results suggest the feeding ecologies of the pygmy and dwarf sperm whales are similar and both species occupy equivalent trophic niches in the region.     

A new species of Isocyamus (Amphipoda: Cyamidae) from Kogia breviceps (De Blainville, 1838) in Australian waters

Isocyamus kogiae n. sp. is described from a pygmy sperm whale Kogia breviceps (De Blainville, 1838) stranded on Moreton Island, Queensland, Australia. It belongs in the genus Isocyamus because of its white appearance, epaulette-like processes at the anterior dorsum of pereon segment 2, lateral spines on segments 3 and 4, reduced maxilliped, and absence of maxilliped palps, large accessory gills and dentate comb on the dactyle of gnathopod 1. It differs from Isocyamus delphini (Guerin-Meneville, 1836), the only other member of this genus, in that the accessory gills are not equal in length to the main gills, it has no ventral spines, and the dactylus of percopod 1 has a double rather than a single dentate comb. This is the first report of a cyamid from whales of the genus Kogia.     

Trypauchen vagina (Bloch & Schneider, 1801) a new established species in the Southwestern Atlantic

Trypauchen vagina (Bloch & Schneider, 1801) is a goby that lives burrowed into the substrata feeding on small invertebrates. It is native to the Indo-pacific region, ranging from Kuwait to China. Recently, this fish has been reported outside the original range of distribution, being found in the Mediterranean Sea, and more recently in the northeastern Brazilian coast. The Mediterranean reports are usually associated with Lessepsian migration, while the reports from Brazil are possibly related to ballast water transportation. In the present work, we provide eight new records from southeastern Brazilian coast, all made in São Paulo state, far from the first record. These additional records raised concern since the presence of an alien species could implicate in environmental and economic losses. Thus, we decide to model the environmental suitability for this goby in the Brazilian coast, specially focusing on major ports, usually places with high ballast water propagule pressure. In addition, an analysis of the suitability in the Red Sea was also made, to verify the hypothesis of Lessepsian migration. The results revealed that temperature and primary productivity are among the most important parameters for the presence of T. vagina, also indicating a high environmental suitability for this species in the Red Sea and Brazilian coast, especially in southeastern region, where the new records were made. Due to the number of collected individuals, it is hard to affirm that this taxon presents a self-sustaining population in Brazilian waters, but the several registers, in different locations and different life stages point to an establishment of the species in this new region. This population status allied with a high environmental suitability is alarming and should motivate new studies concerning T. vagina in Brazilian waters.     

Rod Monochromacy and the Coevolution of Cetacean Retinal Opsins

Cetaceans have a long history of commitment to a fully aquatic lifestyle that extends back to the Eocene. Extant species have evolved a spectacular array of adaptations in conjunction with their deployment into a diverse array of aquatic habitats. Sensory systems are among those that have experienced radical transformations in the evolutionary history of this clade. In the case of vision, previous studies have demonstrated important changes in the genes encoding rod opsin (RH1), short-wavelength sensitive opsin 1 (SWS1), and long-wavelength sensitive opsin (LWS) in selected cetaceans, but have not examined the full complement of opsin genes across the complete range of cetacean families. Here, we report protein-coding sequences for RH1 and both color opsin genes (SWS1, LWS) from representatives of all extant cetacean families. We examine competing hypotheses pertaining to the timing of blue shifts in RH1 relative to SWS1 inactivation in the early history of Cetacea, and we test the hypothesis that some cetaceans are rod monochomats. Molecular evolutionary analyses contradict the “coastal” hypothesis, wherein SWS1 was pseudogenized in the common ancestor of Cetacea, and instead suggest that RH1 was blue-shifted in the common ancestor of Cetacea before SWS1 was independently knocked out in baleen whales (Mysticeti) and in toothed whales (Odontoceti). Further, molecular evidence implies that LWS was inactivated convergently on at least five occasions in Cetacea: (1) Balaenidae (bowhead and right whales), (2) Balaenopteroidea (rorquals plus gray whale), (3) Mesoplodon bidens (Sowerby''s beaked whale), (4) Physeter macrocephalus (giant sperm whale), and (5) Kogia breviceps (pygmy sperm whale). All of these cetaceans are known to dive to depths of at least 100 m where the underwater light field is dim and dominated by blue light. The knockout of both SWS1 and LWS in multiple cetacean lineages renders these taxa rod monochromats, a condition previously unknown among mammalian species.     

The density of odontocete integument depends on blubber lipid composition and temperature

Cetacean integument serves many functional roles, including contribution to whole body buoyancy. The blubber of the integument of different cetacean species contains varying concentrations of triacylglycerols (TAG) and wax esters (WE); generally, these lipid classes have different densities. Integument can also experience a wide range of temperatures during a dive, so its density may change with depth. The goals of this study were to measure integument density and isolated blubber lipid density in three deep‐diving odontocete species (n = 3–4)—short‐finned pilot whales (Globicephala macrorhynchus), pygmy sperm whales (Kogia breviceps), and Gervais' beaked whales (Mesoplodon europeaus)—at different temperatures (6°C–35°C), and to relate these densities to lipid content and composition. Kogia and Mesoplodon integument and isolated lipids had high WE content (78.7–99.5 wt%) and were less dense (by 1.7%–9.3%) than those of Globicephala, which were composed predominately of TAG. Generally, densities increased as temperature decreased. Changes in integument densities mirrored those of isolated lipid densities, suggesting that blubber lipids are largely responsible for the buoyant properties of cetacean integument. These data demonstrate that the contribution of the integument to whole body density depends on lipid class and temperature, and therefore may provide useful, species‐specific correction factors for diving energetics models.     

Temporal variation in dwarf sperm whale (Kogia sima) habitat use and group size off Great Abaco Island, Bahamas

Dwarf sperm whales (Kogia sima) are among the most commonly stranded yet least known pelagic cetaceans. Few studies have occurred at sea, and none have quantified temporal and spatial variation in dwarf sperm whale abundance and group size. We assessed seasonal and spatial variation in dwarf sperm whale group size and abundance off Great Abaco Island, Bahamas using surveys within a 126 km² study area with depths between 2 and 1,600 m. After correcting for survey effort and variation in sighting efficiency among sea states, we found that dwarf sperm whale group size and habitat use varied seasonally. In summer, dwarf sperm whale groups were small (median = 2.5, range = 1–8) and were found only in the two deep habitats within the study area (slope 400–900 m, deep 900–1,600 m). In winter, group sizes increased (median = 4, range = 1–12) and sightings were almost six times higher in the slope habitat, where vertical relief is highest, than other habitats. Our results suggest that studies of pelagic cetaceans and conservation plans must explicitly account for seasonal variation in group size and habitat use.     

Can habitat modelling for the octopus Eledone cirrhosa help identify key areas for Risso’s dolphin in Scottish waters?

Marine protected areas (MPAs) are becoming increasingly important tools in the conservation of cetaceans. This has led to an interest in the use of species distribution models to predict where cetacean species are likely to occur based on the distribution of environmental variables. However, relationships between cetacean distribution and environmental variables are generally assumed to reflect the environmental preferences of their prey. Thus, understanding the distribution of prey may increase our ability to identify important areas for cetaceans. Here, we describe the diet of Risso’s dolphin (Grampus griseus) by analysing stomach contents of individuals stranded in Scotland over the last twenty years. Next, we use habitat modelling to test whether Rissos’s dolphin distribution in Scottish waters, as inferred from sightings, is related to the distribution of its main prey, the octopus Eledone cirrhosa. While good models of the relationship between the distribution of E. cirrhosa and environmental variables were obtained, there was no evidence of a relationship between modelled octopus distribution and the occurrence of Risso’s dolphins. These results suggest that identifying key areas for its main prey species is unlikely to help identify potential MPAs for Risso’s dolphin, at least at the spatial resolution used in this study.     

Occurrence of hourglass dolphin (Lagenorhynchus cruciger) and habitat characteristics along the Patagonian Shelf and the Atlantic Ocean sector of the Southern Ocean

The hourglass dolphin, Lagenorhynchus cruciger, is one of the smallest cetaceans found in Antarctic waters. Although frequently observed, this is one of the least studied species of dolphin. In this study, we investigate the occurrence of hourglass dolphin in the South-West Atlantic and Southern Ocean from 42°S to 63°S and characterize the oceanographic and environmental features associated with their occurrence. Sighting data were collected during two scientific cruises in December 2009 and January and February 2011 and integrated into a geographical information system with environmental variables such as sea surface temperature, ocean colour, depth, distance to oceanographic fronts and distance from shore. We used a principal component analysis (PCA) to identify patterns in data and a non-hierarchic cluster analysis (K means) to classify observations into groups. A total of 19 sightings (96 individuals) were recorded. Group size ranged from one to twelve dolphins (mean 5.05). We could differentiate two groups in relation to the environmental conditions where animals were found, one group occurring in shallow coastal waters and the other occurring in deeper and colder waters further offshore. The present study contributes novel information on environmental measures associated with hourglass dolphin distribution in the South-West Atlantic and Southern Oceans. Additionally, it includes relatively high sighting clusters of this species in two highly productive and biodiverse areas (Burdwood Bank/Isla de los Estados and the South Shetland Islands).