A new deep-sea hydroid (Cnidaria: Hydrozoa) from the Bering Sea Basin reveals high genetic relevance to Arctic and adjacent shallow-water species

The marine benthic fauna in Arctic shallow-water is reported to be a relatively young assemblage by species of either Pacific or Atlantic affinity. Whether current deep-sea Pacific species are included in the affinity or not is unknown. Combining morphological comparisons and genetic analyses, a new deep-sea hydroid to science, Sertularia xuelongi sp. nov. (Cnidaria: Hydrozoa: Sertulariidae), is described from the northern margin of the Bering Sea Basin at depths of 800–1570 m collected in 2010. It is characterized by slender and zigzag-shaped hydrocauli, alternately arranged hydrothecae and the absence of distal-lateral horns in fully matured female gonothecae. Its distribution, currently known only from Bering Sea Basin, suggests that it could not be an Arctic species with Pacific affinity. However, phylogenetic analyses based on the mitochondrial 16S rRNA gene show that it is clustered into a distinctive clade with four closely related species recorded from shallow-water of Northwest France, Iceland, Chukchi Sea and/or Bering Sea. In addition, its sequence similarity is highly relevant to these four species: Sertularia argentea (98.6 %), S. cupressina (98.8 %), S. plumosa (98.8 %) and S. robusta (99.4 %). All these provide a new insight into the relevance of North Pacific deep-sea species to the benthic fauna in Arctic and adjacent shallow-water. The taxonomic restriction of the genus Sertularia and the re-validation of the genus Polyserias are discussed. Future researches on more deep-sea species from Pacific and/or Atlantic are required to understand the evolution and speciation pattern involved in polar relevance.     

Macrofaunal crustaceans in the benthic boundary layer from the shelf break to abyssal depths in the Ross Sea (Antarctica)

The New Zealand International Polar Year–Census of Antarctic Marine Life (NZ IPY-CAML) project added to previous benthic studies in the Ross Sea by extending sampling north from the continental shelf to previously unsampled areas of the shelf break, slope, abyssal plain and seamounts in the region. The aim of the current study is to give first insights into the deep-sea community structure of the Ross Sea focussing on a component of the benthic boundary layer that is macrofaunal crustaceans collected one metre above the seafloor. We assess changes in Ross Sea crustacean community composition from the shelf break (474 m) to the abyss (3,490 m) and compare the Ross Sea crustacean fauna to areas elsewhere in the Southern Ocean. Analyses reveal high relative abundances, suggesting an important role in the food web. Among the peracarid crustaceans, there was a decline in the proportion of amphipods with increasing depth. Three of 15 isopod families (Acanthaspidiidae, Nannoniscidae and Desmosomatidae) were identified to species level and about 72 % of the species were new to science. Isopod diversity in the Ross Sea abyss appears to be comparable to that in the highly speciose Weddell deep sea. Standardised sampling of these crustacean communities allows setting the biodiversity of the Ross Sea into a global context.     

Mesozooplankton distribution patterns and grazing impacts of copepods and Euphausia crystallorophias in the Amundsen Sea, West Antarctica, during austral summer

The rapid melting of glaciers as well as the loss of sea ice in the Amundsen Sea makes it an ideal environmental setting for the investigation of the impacts of climate change in the Antarctic on the distribution and production of mesozooplankton. We examined the latitudinal distribution of mesozooplankton and their grazing impacts on phytoplankton in the Amundsen Sea during the early austral summer from December 27, 2010 to January 13, 2011. Mesozooplankton followed a latitudinal distribution in relation to hydrographic and environmental features, with copepods dominating in the oceanic area and euphausiids dominating in the polynya. Greater Euphausia crystallorophias biomass in the polynya was associated with lower salinity and higher food concentration (chlorophyll a, choanoflagellates, and heterotrophic dinoflagellates). The grazing impact of three copepods (Rhincalanus gigas, Calanoides acutus, and Metridia gerlachei) on phytoplankton was low, with the consumption of 3 % of phytoplankton standing stock and about 4 % of daily primary production. Estimated daily carbon rations for each of the three copepods were also relatively low (<10 %), barely enough to cover metabolic demands. This suggests that copepods may rely on food other than phytoplankton and that much of the primary production is channeled through microzooplankton. Daily carbon rations for E. crystallorophias were high (up to 49 %) with the grazing impact accounting for 17 % of the phytoplankton biomass and 84 % of primary production. The presence of E. crystallorophias appears to be a critical factor regulating phytoplankton blooms and determining the fate of fixed carbon in the coastal polynyas of the Amundsen Sea.     

<Emphasis Type="Italic">Campylonotus arntzianus</Emphasis>, a new species of the Campylonotidae (Crustacea: Decapoda: Caridea) from the Scotia Sea (Antarctica)

Two specimens of Campylonotus arntzianus sp. nov. were caught in the Antarctic Scotia Sea off Saunders Island (57°40.31'S, 26°27.81'W) using an Agassiz trawl at one station (depth: 475-589 m). The new species described here is the fifth representative of the monogeneric family Campylonotidae, and the first of the family south of the Antarctic Convergence. Campylonotus arntzianus sp. nov. is a shrimp of about 5 cm in total length. Due to similarities in adult morphology, C. arntzianus sp. nov. seems to be closely related to C. capensis, a deep-sea species from the Southern Atlantic Ocean. A simple key for the species identification of the Campylonotidae is provided.     

The trophodynamics of Southern Ocean Electrona (Myctophidae) in the Scotia Sea

The Scotia Sea is one of the most productive regions of the Southern Ocean, but its surface waters are experiencing a rapid increase in temperature, which may be changing the behaviour and distribution of many myctophids and their prey species. Electrona antarctica and Electrona carlsbergi are two of the most abundant myctophids in the region, but their ecology is poorly understood and their response to ongoing environmental change is difficult to determine. This study investigated spatial and temporal patterns in their abundance, population structure and diets using mid-water trawl nets deployed across the Scotia Sea during spring, summer and autumn. E. antarctica was the most numerically abundant species (0.09–0.21 ind. 1,000 m^?3), with greatest concentrations occurring in the sea-ice sectors. E. carlsbergi occurred in more northern regions, comprising densities of 0.02–0.11 ind. 1,000 m^?3. There was evidence of seasonal variation in depth distribution, size-related sexual dimorphism and size-specific vertical stratification for both species. Latitudinal trends in sex ratio and female body size were apparent for E. antarctica. Its diet varied between regions, seasons and size classes, but overall, Euphausia superba, Metridia spp. and Themisto gaudichaudii were the dominant prey items. E. carlsbergi appeared not to recruit in the Scotia Sea. Its diet was dominated by copepods, particularly Rhincalanus gigas and Metridia spp., but regional, seasonal and ontogenetic variations were evident. This study contributes to our understanding of how mid-water food webs are structured in the Southern Ocean and their sensitivity to ongoing environmental change.     

Larvae of the deep-sea Nematocarcinidae (Crustacea: Decapoda: Caridea) from the Southern Ocean

The early larval stages of the deep-sea Nematocarcinidae, Nematocarcinus longirostris Bate, 1888, from the south-western Atlantic Ocean, and N. lanceopes Bate, 1888, from the high Antarctic Weddell Sea, were obtained from plankton catches, described and illustrated. Furthermore, field collected larvae of N. lanceopes were compared with larvae hatched and reared under constant laboratory conditions. The morphology of larvae in both species clearly indicates a planktotrophic and extended mode of larval development. This is an outstanding feature in deep-sea and especially in high-latitudinal caridean shrimp species, and the consequence of such reproductive trait for life history adaptations to both deep-sea and polar environments is discussed.This revised version was published online in November 2004 with corrections to the figure part lettering.     

Spatial distribution of phytoplankton productivity in the Amundsen Sea, Antarctica

To date, no direct measurements of primary production were taken in the Amundsen Sea, which is one of the highest primary productivity regions in the Antarctic. Phytoplankton carbon and nitrogen uptake experiments were conducted at 16 selected stations using a ¹³C–¹⁵N dual isotope tracer technique. We found no statistically significant depletions of major inorganic nutrients (nitrate?+?nitrite, ammonium, and silicate) although the concentrations of these nutrients were markedly reduced in the surface layer of the polynya stations where large celled phytoplankton (>20?μm) predominated (ca. 64?%). The average chl-a concentration was significantly higher at polynya stations than at non-polynya stations (p?<?0.01). Average daily carbon and nitrogen uptake rates by phytoplankton at polynya stations were 2.2?g?C?m^?2?day^?1 (SD?=?±1.4?g?C?m^?2?day^?1) and 0.9?g?N?m^?2?day^?1 (SD?=?±0.2?g?N?m^?2?day^?1), respectively, about 5–10 times higher than those at non-polynya stations. These ranges are as high as those in the Ross Sea, which has the highest productivity among polynyas in the Antarctic Ocean. The unique productivity patterns in the Amundsen Sea are likely due to differences in iron limitation, phytoplankton productivity, the timing of phytoplankton growing season, or a combination of these factors.     

Upper Eocene to Middle Miocene ostracode faunas and paleo-oceanography of the North Coastal Belt,Jamaica, West Indies

The ostracode fauna of the Montpelier Group (Upper Eocene-Middle Miocene) exposed in the western part of the North Coastal Belt of Jamaica contains representatives of three paleo-environments. Allochthonous shelf and littoral species, characterized by abraded carapaces and eye tubercles, and a diverse group of archibenthal (slope) forms are present throughout. Species restricted to the World Ocean Psychrosphere (> 1000 m), notably Bradleya dictyon, Australoecia tipica, Agrenocythere hazelae and Macrocypris spp., first occur in sediments of earliest Miocene age (planktonic foraminiferal zone N4). Their presence establishes a minimum age for the entry into the Cayman Trench of frigid (<8–10°C) water masses drawn from the Atlantic thermohaline stratification.Upper Eocene and Oligocene assemblages are dominated by smooth, blind genera such as Krithe, Messinella and Bythocypris and lived in lower thermospheric conditions (10°C) at or near the base of the thermocline (700– 900 m). During the Early Miocene, downfaulting of the north island slope of Jamaica in response to southward tilting of the Nicaraguan Rise brought the Montpelier depositional site into contact with progressively deeper and colder water-masses. This subsidence culminated in the late Early Miocene; faunas of this age contain large numbers of Bradleya dictyon, Agrenocythere hazelae, Macrocypris sp. 1, Bairdia oarion and Bairdoppilata cassida. They suggest paleodepths of 1500–2000 m and the local presence of a 3–4° water-mass tentatively identified as the North Atlantic deep water. If this interpretation is correct, then the maximum sill-depth and water-mass configuration of the Miocene Cayman Trench have persisted to the present. Middle Miocene assemblages indicate a return to paleodepths of 1000–1500 m heralding the pronounced Upper Miocene to mid-Pleistocene tectonism which elevated Montpelier strata above sea level. An alternative view attributes the Miocene ostracode succession in the Montpelier to episodes in the development of the Atlantic circulation as recognized in the northeast Atlantic.Montpelier deposition took place either on a broad step intervening between the northern edge of the Clarendon Platform and the abyssal basins of the Cayman Trench or on a Tertiary analog of the existing north island slope. These results have emphasized the value of detailed taxonomic and morphologic studies of deep-sea ostracodes in geohistorical reconstruction.     

Pre-winter distribution and habitat characteristics of polar cod (Boreogadus saida) in southeastern Beaufort Sea

Polar cod was shown to form dense under-ice winter aggregations at depth in the Amundsen Gulf (southeastern Beaufort Sea). In this paper, we verify the premises of the aggregation mechanism by determining the distribution and habitat characteristics of polar cod prior to the formation of winter aggregations. Multifrequency split-beam acoustic data collected in October–November 2003 revealed that polar cod split into two distinct layers. Age-0 polar cod formed an epipelagic layer between 0 and ~60 m depth without any clear large-scale biomass trend. In contrast, adult polar cod tended to distribute into an offshore mesopelagic layer between ~200 and 400 m that shoaled into a denser (1–37 g m^?2) benthopelagic layer on sloping bottoms (between 150 and 600-m isobaths) along the Mackenzie shelf and into the Amundsen Gulf basin. Concentrations peaked in the Amundsen Gulf where estimated total biomass reached ~250 kt. Both age-0 and adult polar cod distributed in the warmer waters (>?1.4 °C). We hypothesise that polar cod concentration over slopes is governed by the combined actions of (1) local currents concentrating both depth-keeping zooplankton and polar cod at the shelf-break and basin slopes and (2) trophic association with these predictable topographically trapped aggregations of zooplankton prey. During freeze-up, these slope concentrations of polar cod are thought to constitute the main source of the observed dense under-ice winter aggregations. The hypothesis of active short-distance displacements combined with prevailing mean currents is retained as the likely aggregation mechanism.     

Catching of Greenland halibut <Emphasis Type="Italic">Reinhardtius hippoglossoides</Emphasis> (Pleuronectidae) on the shelf edge of the Laptev and East Siberian Seas

The first capture of the Greenland (or Black) halibut Reinhardtius hippoglossoides in the Arctic Ocean on the border with the East Siberian Sea (79°03′–79°08′ N 139°59′–141°16′ E, 259–277 m) and the repeated findings of the species in the Laptev Sea (78°03′–78°04′ N 132°56′–133°04′ E, 307 m; 78°33′–78°35′ N 138°44′–138°48′ E, 125 m) are reported. Fish (79 specimens with the length of 15.0–44.5 cm) were caught during four bottom trawlings on the edge of the continental shelf of the two seas in the transformed waters of Atlantic origin, as evidenced by the discovery of the indicator species of these waters—the Glacier lanternfish Benthosema glaciale (Myctophidae)—in one of the stomachs of halibut. The other captures of Greenland halibut in the Arctic were also reviewed.     

A new species of Wellsopsyllus (Copepoda, Harpacticoida, Paramesochridae) from the deep Southern Ocean and remarks on its biogeography

The new deep-sea copepod species of the family Paramesochridae (Copepoda, Harpacticoida) presented here was collected during the scientific ANtarctic benthic DEEP-sea biodiversity cruise II (ANDEEP II) to the Weddell Abyssal Plain and western Weddell Sea and the CROZet natural iron bloom and EXport experiment (CROZEX cruise) to the Crozet Isles in the Indian Ocean. The new species has been allocated to the subgenus Wellsopsyllus within the genus Wellsopsyllus Kunz, 1981, based on the 1-segmented endopods and 3-segmented exopods of swimming legs 2–4. Furthermore, the new species shows a 1-segmented exopod of the antenna. Wellsopsyllus (W.) antarcticus sp. n. can be distinguished from its congeners by its small body size, the presence of two setae on the distal segment of the endopod of the first swimming leg, by the very small furcal rami, the extremely reduced endopods of second and third swimming legs, the reduced mandibular palp and maxillula, as well as by the size and armature of the fifth and sixth swimming legs in both sexes. Difficulties of the placement of the new species into a suitable genus show an insufficiency in the present family classification. In the future, a revision of the genus Wellsopsyllus is essential. With its presence in the western Weddell Sea and the Indian Ocean, the new species may have a wide distribution range. Furthermore, it is the first abyssal species of the genus Wellsopsyllus sampled outside of the Scotia Sea.     

Under-ice distribution of polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> in the central Arctic Ocean and their association with sea-ice habitat properties

In the Arctic Ocean, sea-ice habitats are undergoing rapid environmental change. Polar cod (Boreogadus saida) is the most abundant fish known to reside under the pack-ice. The under-ice distribution, association with sea-ice habitat properties and origins of polar cod in the central Arctic Ocean, however, are largely unknown. During the RV Polarstern expedition ARK XXVII/3 in the Eurasian Basin in 2012, we used for the first time in Arctic waters a Surface and Under Ice Trawl with an integrated bio-environmental sensor array. Polar cod was ubiquitous throughout the Eurasian Basin with a median abundance of 5000 ind. km^?2. The under-ice population consisted of young specimens with a total length between 52 and 140 mm, dominated by 1-year-old fish. Higher fish abundance was associated with thicker ice, higher ice coverage and lower surface salinity, or with higher densities of the ice-amphipod Apherusa glacialis. The fish were in good condition and well fed according to various indices. Back-tracking of the sea-ice indicated that sea-ice sampled in the Amundsen Basin originated from the Laptev Sea coast, while sea-ice sampled in the Nansen Basin originated from the Kara Sea. Assuming that fish were following the ice drift, this suggests that under-ice polar cod distribution in the Eurasian Basin is dependent on the coastal populations where the sea-ice originates. The omnipresence of polar cod in the Eurasian Basin, in a good body condition, suggests that the central Arctic under-ice habitats may constitute a favourable environment for this species survival, a potential vector of genetic exchange and a recruitment source for coastal populations around the Arctic Ocean.     

Is the Scotia Sea a centre of Antarctic marine diversification? Some evidence of cryptic speciation in the circum-Antarctic bivalve <Emphasis Type="Italic">Lissarca notorcadensis</Emphasis> (Arcoidea: Philobryidae)

The bivalve Lissarca notorcadensis is one of the most abundant species in Antarctic waters and has colonised the entire Antarctic shelf and Scotia Sea Islands. Its brooding reproduction, low dispersal capabilities and epizoic lifestyle predict limited gene flow between geographically isolated populations. Relationships between specimens from seven regions in the Southern Ocean and outgroups were assessed with nuclear 28S rDNA and mitochondrial cytochrome oxidase subunit I (COI) genes. The 28S dataset indicate that while Lissarca appears to be a monophyletic genus, there is polyphyly between the Limopsidae and Philobryidae. Thirteen CO1 haplotypes were found, mostly unique to the sample regions, and two distinct lineages were distinguished. Specimens from the Weddell and Ross Sea form one lineage while individuals from the banks and islands of the Scotia Sea form the other. Within each lineage, further vicariance was observed forming six regionally isolated groups. Our results provide initial evidence for reproductively isolated populations of L. notorcadensis. The islands of the Scotia Sea appear to act as centres of speciation in the Southern Ocean.     

Conquered from the Deep Sea? A New Deep-Sea Isopod Species from the Antarctic Shelf Shows Pattern of Recent Colonization

The Amundsen Sea, Antarctica, is amongst the most rapidly changing environments of the world. Its benthic inhabitants are barely known and the BIOPEARL 2 project was one of the first to biologically explore this region. Collected during this expedition, Macrostylis roaldi sp. nov. is described as the first isopod discovered on the Amundsen-Sea shelf. Amongst many characteristic features, the most obvious characters unique for M. roaldi are the rather short pleotelson and short operculum as well as the trapezoid shape of the pleotelson in adult males. We used DNA barcodes (COI) and additional mitochondrial markers (12S, 16S) to reciprocally illuminate morphological results and nucleotide variability. In contrast to many other deep-sea isopods, this species is common and shows a wide distribution. Its range spreads from Pine Island Bay at inner shelf right to the shelf break and across 1,000 m bathymetrically. Its gene pool is homogenized across space and depth. This is indicative for a genetic bottleneck or a recent colonization history. Our results suggest further that migratory or dispersal capabilities of some species of brooding macrobenthos have been underestimated. This might be relevant for the species’ potential to cope with effects of climate change. To determine where this species could have survived the last glacial period, alternative refuge possibilities are discussed.     

Palaeoenvironmental significance of Miocene larger benthic foraminifera from the Xisha Islands,South China Sea

We studied 344 samples from Well XK-1 in Xisha Islands, South China Sea, and identified 66 species of larger benthic foraminifera, providing critical evidence for biostratigraphy and palaeoenvironmental interpretation of the Miocene reef carbonate sequence. Three assemblages are recognized, namely, Spiroclypeus higginsi–Borelis pygmaeus Assemblage (Letter Stage Te5, Early Miocene, 1256.28–1180.15 m), Nephrolepidina–Miogypsina Assemblage (Tf, Middle Miocene, 1031.10–577.04 m), and Cycloclypeus–Heterostegina Assemblage (Tg, Late Miocene, 468.13–380.42 m). On the basis of the palaeoecological preference of the larger foraminifera, we interpret that the Miocene carbonate sequence was deposited mainly in a warm tropical shallow water environment, characterized by five stages of continuous long-term evolution: backreef lagoon to shelf in the Early Miocene, normal to frontal reef in the early Middle Miocene, backreef lagoon to shelf in the later Middle Miocene, normal to frontal reef in the early Late Miocene, and proximal forereef shelf in the later Late Miocene.     

Species diversity and distribution within the deep-sea nematode genus Acantholaimus on the continental shelf and slope in Antarctica

Species diversity and distribution—in relation with biogeography, bathymetry and environmental characteristics—within the genus Acantholaimus were investigated on the shelf and slope (182–2,009 m) in the Weddell Sea (Kapp Norvegia and Vestkapp) and near the Antarctic Peninsula (Drake Passage and Bransfield Strait). The occurrence of the otherwise typical deep-sea genus Acantholaimus in high densities and diversity on the continental shelf is a unique feature of the Southern Ocean. Fifty-five Acantholaimus species (morpho-types) were distinguished in our study area, of which 39 were found on the shelf. The Acantholaimus communities were characterised by a high number of congeneric species and the presence of many rare species, similar to typical deep-sea communities. Fourteen species had a distribution extending from the shelf to the lower slope. High local and regional diversity was obvious and a considerable level of endemism was suggested. A negative impact of chlorophyll a concentrations, reflecting the availability of phytoplankton, was found on the Acantholaimus densities and species diversity.     

Life history strategies of the Scotia Sea icefish, <Emphasis Type="Italic">Chaenocephalus aceratus</Emphasis>, along the Southern Scotia Ridge

Reproductive capacity can influence distribution and abundance over large spatial scales through larval dispersal, even when adult stages remain isolated following settlement. We examined size distribution, reproductive traits and age structure in Scotia Sea icefish, Chaenocephalus aceratus, an abundant benthic species with a long larval pelagic phase found on continental shelves along the Southern Scotia Ridge. In particular, we compared life history strategies between fish caught during surveys undertaken off the South Orkney Islands (SOI) and South Shetland Islands (SSI). Results corroborated regional separation after settlement and suggested distinct life history strategies, in which fish from SOI invested much less in reproduction, and somewhat more in somatic growth earlier in their life history. Compared to SSI, body weight increased faster with length and absolute fecundity was 46 % lower and increased more slowly with size for SOI population. In addition, the proportion of spawning cohorts and L _∞ was lower and k higher for SOI. The differences appeared to be a phenotypic response to environmental conditions related to regional hydrography. Lower reproductive capacity around the SOI, and strong eastward flow in the large-scale circulation, suggests that the SSI may be more important in influencing distributions and abundance of icefish along the Southern Scotia Ridge.     

Summer social structure of crabeater seal <Emphasis Type="Italic">Lobodon carcinophaga</Emphasis> in the Amundsen Sea,Antarctica

During the PS75 expedition of ice-breaking RV Polarstern in the Amundsen Sea, in February–March 2010, we studied the at-sea quantitative distribution of the “upper trophic levels,” seabirds and marine mammals. In the Amundsen Sea Embayment, 14,200 pinnipeds belonging to four species were recorded (2100 during 670 half-hour transect counts from the bridge and 12,100 during 50 h of helicopter flight). Crabeater seal Lobodon carcinophaga represented more than 97 % of the total. Two types of major aggregations of crabeaters were noted: on the one hand, very large groups hauled out on the pack ice, composed mainly of adults/bachelors and very few calves. On the other hand, swimming pods of calves were accompanied by one or two leading adults around icebergs. Our interpretation is that, after the beginning of the reproduction period with trios (cow, calf and future father) scattered on pack ice, such pods can be considered the last step of the breeding cycle.     

An unusual hermaphrodite reproductive trait in the Antarctic brooding bivalve Lissarca miliaris (Philobryidae) from the Scotia Sea, Southern Ocean

The Antarctic marine environment is extreme in its low temperatures and short periods of primary productivity. Invertebrates must therefore adapt to maximise reproductive output where low temperature and limited food slow larval development. Brooding is a common reproductive trait in Antarctic marine bivalves; larval development occurs within the mantle cavity, and larvae are released as fully developed young. Lissarca miliaris is a small, short-lived, shallow-water brooding bivalve of circum-Antarctic distribution and found most abundant in the sub-Antarctic Magellan Region and islands of the Scotia Arc. Here, an unusual hermaphrodite reproductive trait is described for L. miliaris from King George Island (62°14′S, 58°38′W) and Signy Island (60°42′S, 45°36′W), Antarctica, using histological and dissection techniques. Specimens demonstrate simultaneous and sequential hermaphrodite traits; male and female gonads develop simultaneously, but the production of oocytes is reduced while testes are ripe. Functional females are more abundant in specimens above 3 mm shell length, although male reproductive tissue persists and functional males are found in all size classes. The number of previtellogenic oocytes produced by far exceeds the number of oocytes extruded and brooded, which may indicate an ancestral link to a planktotrophic past. Hermaphroditism in L. miliaris maximises reproductive efficiency in a short-lived species, in which the female’s capacity to brood its young is limited, and demonstrates a specialised adaptation to a cold stenothermal and food-limited environment prevailing in the Southern Ocean.