Demographic history and population structure of the Antarctic silverfish Pleuragramma antarcticum

The Antarctic silverfish Pleuragramma antarcticum (Nototheniidae) is the most abundant pelagic fish inhabiting Antarctic waters. In this study we investigated, through partial sequencing of the D-loop mitochondrial region, samples collected at four different locations in the Southern Ocean, three in the Atlantic and one in the Pacific sector. Sampling was replicated in two different years at two locations. Sequence analysis showed a remarkably high polymorphism, with 110 haplotypes over the 256 investigated specimens, and about 80% of haplotypes occurring only once. Neutrality tests indicated that all samples were not at mutation-drift equilibrium, and suggested a past population expansion. This result was supported by the presence of a star-like topology in the D-loop gene tree, and by results of mismatch distribution. The start of the expansion was dated, using a specifically calibrated clock, between 111 and 126 thousand years ago. This value corresponds to the start of the cooling period that led to the last glaciation peak, and is in close agreement with a recently suggested range expansion for pelagic Antarctic ecosystems. Analysis of molecular variation indicated a small, though highly significant, value of differentiation between samples. This result, together with the lack of association between clades and geographical locations, indicates a weak population structure for the species.     

Metabolic and behavioural adaptations during early development of the Antarctic silverfish, Pleuragramma antarcticum

The Antarctic silverfish Pleuragramma antarcticum is a keystone species in the Ross Sea ecosystem, providing one of the major links between lower and higher trophic levels. Despite the importance of this species, surprisingly little is known of its early development and behaviour. Here, we determine the metabolic capacity of Pleuragramma embryonated eggs and larvae and make comparisons with developing stages of another notothenioid, the naked dragonfish Gymnodraco acuticeps. We also show that although large numbers of embryonated eggs of Pleuragramma are found floating among the platelet ice of Terra Nova Bay, they are able to sink prior to hatching in late spring, likely reducing the risk of exposure to the potentially lethal, ice-laden surface environment. Applying Stoke’s law, we determine the change in density required for embryonated eggs to sink at the measured rate and then consider possible mechanisms by which this might occur. Significantly, newly hatched larvae are positively gravitactic and negatively phototactic, such that their swimming behaviour also directs them away from the risk of freezing in the icy surface waters. Measurement of the acute thermal tolerance shows that Pleuragramma larvae have, on average, a sustainable swimming performance breadth of about 17°C, which is significantly greater than that of other adult notothenioids. Although it lacks significant antifreeze capacity in its early developmental stages, Pleuragramma has other attributes that may ensure survival over a wider range of environmental temperatures than other more stenothermal Antarctic notothenioids. How it might adapt to prolonged environmental change arising from phenomena such as global warming, however, requires further investigation.     

Diet and trophic niche of Antarctic silverfish Pleuragramma antarcticum in the Ross Sea,Antarctica

The diet of Antarctic silverfish Pleuragramma antarcticum was evaluated by examining stomach contents of specimens collected in the Ross Sea (71°–77° S; 165°–180° E) in January to March 2008. Pleuragramma antarcticum (50–236 mm standard length, L_S) and prey items were analysed for stable‐isotopic composition of carbon and nitrogen. According to index of relative importance (I_RI), which incorporates frequency of occurrence, mass and number of prey items, the most important prey items were copepods (81%I_RI over all specimens), predominantly Metridia gerlachei and Paraeuchaeta sp., with krill and fishes having low I_RI (2·2 and 5·6%I_RI overall). According to mass of prey (M) in stomachs, however, fishes (P. antarcticum and myctophids) and krill dominated overall diet (48 and 22%M, respectively), with copepods being a relatively minor constituent of overall diet by mass (9·9%M). Piscivory by P. antarcticum occurred mainly in the extreme south‐west of the region and near the continental slope. Krill identified to species level in P. antarcticum stomachs were predominantly Euphausia superba (14·1%M) with some Euphausia crystallophorias (4·8%M). Both DistLM modelling (PRIMER‐permanova+) on stomach contents (by I_RI) and stepwise generalized linear modelling on stable isotopes showed that L_S and location were significant predictors of P. antarcticum diet. Postlarval P. antarcticum (50–89 mm L_S) consumed exclusively copepods. Juvenile P. antarcticum (90–151 mm L_S) consumed predominantly krill and copepods by mass (46 and 30%M, respectively). Small adult P. antarcticum (152–178 mm L_S) consumed krill, fishes and copepods (37, 36 and 15%M, respectively). Large adult P. antarcticum (179–236 mm L_S) consumed predominantly fishes and krill (55 and 17%M, respectively), especially in the north (near the Ross Sea slope) and in the SW Ross Sea. Amphipods were occasionally important prey items for P. antarcticum (western Ross Sea, 39%M). General concordance between stomach contents and trophic level of P. antarcticum and prey based on δ¹⁵N was demonstrated. Pleuragramma antarcticum trophic level was estimated as 3·7 (postlarval fish) and 4·1 (fish aged 3+ years).     

Sensory development of the Antarctic silverfish Pleuragramma antarcticum: a test for the ontogenetic shift hypothesis

In 1996 Montgomery proposed an ontogenetic shift in the use of visual and non-visual senses in Antarctic notothenioid fishes, with visual dominance in larval fishes giving way to non-visual senses in adults. One prediction of the hypothesis is timing differences in the development of the respective sensory systems, with the visual system expected to develop earlier than the other systems. The volume of certain brain centres can be determined from fixed material and should correlate with sensory development. This study determined the relative volumes of visual and lateral line brain areas, and relative eye size as a function of fish length in Pleuragramma antarcticum.The relative volume of optic tectum was largest in larval fish, exhibiting a negative allometry with growth. The eminentia granularis, and crista cerebellaris (lateral line associated areas) were not recognisable in the smallest larvae; they became differentiated at standard lengths of 10–20 mm and their relative volumes continued to increase over the size range of fish studied (up to 150 mm standard length). Relative eye diameter decreased dramatically over the size range 5–25 mm and then increased such that relative eye diameter doubled over the size range 25–30 mm. A similar, but less extreme, pattern was seen over the size range 30–60 mm. Above 60 mm relative eye diameter increased slightly with size. Our interpretation is that eye growth and somatic growth are on separate trajectories, and the breaks in the relative eye diameter curve result from overwinter periods when somatic growth is static, but the eye continues to grow. These results provide support for the ontogenetic shift hypothesis, and indicate that the timing of the shift probably occurs after the second winter. Received: 22 October 1996 / Accepted: 10 January 1997     

Born among the ice: first morphological observations on two developmental stages of the Antarctic silverfish Pleuragramma antarcticum, a key species of the Southern Ocean

The Antarctic silverfish Pleuragramma antarcticum is a keystone species in the Southern Ocean ecosystem, providing one of the major links between lower and higher trophic levels. Despite the importance of this species, surprisingly little is known of its early development. The first spawning area for the silverfish has been recently identified in the near-shore of Terra Nova Bay (Ross Sea). Evidence indicates that spawning and embryo development occurs in the cryopelagic environment, below the seasonal pack-ice. In order to contribute to the knowledge of the life cycle of this very important Antarctic species, we carried out the first histological characterization on pre-hatching embryos and newly hatched larvae. Embryonated eggs and larvae of P. antarcticum were collected between late October and November 2005 at TNB through holes drilled into the sea ice. Embryonic stage just before hatching and the first post-hatching stage were the most abundant within our samples and thus were analysed using both macroscopic and histological approaches. Early life stages of the Antarctic silverfish revealed interesting features: the sensory system, foraging apparatus and heart appeared well developed, whereas the liver and gills were underdeveloped. Morphological details of the organogenesis were performed, providing the first substantial information on the development of P. antarcticum and representing a further steps towards the knowledge of the life cycle of this important Antarctic key species. An erratum to this article can be found at     

Stomach contents of the Antarctic Silverfish Pleuragramma antarcticum from the southern and eastern Weddell Sea (Antarctica)

Summary The stomach contents of 142 Pleuragramma antarcticum from the southernmost part of the Weddell Sea (Gould Bay: 77°19S) and from the eastern coast (72°–74°S) were analyzed. The fish were collected in February 1983 and stored frozen. Size range of the investigated specimens was from 6.4 cm SL (7.3 cm TL) to 21.9 cm SL (24.1 cm TL). The fish were caught by bottom trawl (Gould Bay) and Agassiz Trawl and pelagic RMT-m net in the upper 300 m layer at the eastern coast, respectively. Thirty-six stomachs were empty or contained sand only. The most important prey in terms of biomass were euphausiids, which occurred in 49 stomachs at a mean number of 2. Their dry weight was estimated to be 15–50 times that of the next important food items, which were copepods, gastropods and gammarids. Krill (Euphausia superba) constituted up to 14% of the euphausiids in the diet of fish caught in the eastern Weddell Sea. The rest was usually made up by Euphausia crystallorophias. This species was taken with preference even when the abundance of E. superba in accompanying plankton catches was 16 times higher. In the Gould Bay, gastropods were frequently eaten despite high numbers of copepods in the plankton, whereas in the eastern Weddell Sea, copepods were abundant both in plankton and in the stomach content. The almost complete absence of the early stages of fish in the diet of Pleuragramma antarcticum in the southern and eastern Weddell Sea is due to a pronounced vertical segregation of fish of different sizes. This distribution pattern is thought to be an adaptive mechanism to avoid intraspecific predation, as 90% of the ichthyoplankton in that area is Pleuragramma antarcticum.     

Geographical distribution and abundance of postlarval and juvenile Pleuragramma antarcticum (Pisces,Notothenioidei) off the Antarctic Peninsula

Summary The early life history of Pleuragramma antarcticum off the Antarctic Peninsula is described from extensive material collected during four expeditions between 1975/76 and 1982 by means of an RMT 1+8. Postlarvae occurred frequently on the northern continental shelf in February and March except during the 1977/78 season when they were almost absent. Abundance varied mainly between 1.8 and 14.7 postlarvae per 10³ m³ with maxima between 32.6 and 143.3 per 10³ m³. Juveniles were less abundant with 0.1–4.9 and maxima between 9.8 and 28.3 per 10³ m³, but formed a consistent, predominant proportion (83–97%) of the pelagic juvenile fish fauna throughout the years of the study. They were distributed over the whole Bransfield Strait and adjacent waters in the top 200 m. Postlarvae and juveniles showed different patterns of vertical distribution; more than 70% of postlarvae occurred in the top 135 m of the water column whereas 83% of juveniles were encountered between 70 and 200 m with increasing abundance towards the lower depth. Three length groups were identified corresponding to age groups 0, 1, and 2. Hatching was estimated to occur in December. Postlarval distribution patterns and the prevailing transport of water masses suggest that coastal waters south of the tip of the Antarctic Peninsula and in the north-eastern Bellingshausen Sea are probable spawning areas. The Bransfield Strait and adjacent waters represent nursery grounds for the early stages of P. antarcticum from these sites.     

A nursery area for the Antarctic silverfish Pleuragramma antarcticum at Terra Nova Bay (Ross Sea): first estimate of distribution and abundance of eggs and larvae under the seasonal sea-ice

Pleuragramma antarcticum is the dominant pelagic fish in the waters of the continental shelf in high Antarctic regions, where it plays a key role in the food web. A nursery ground for eggs of this species was first identified in 2002 in Terra Nova Bay (Ross Sea), where eggs were found trapped in ice platelets under the sea-ice during the spring. As part of a monitoring program aimed at understanding the geographic and temporal characteristics of this nursery ground, the present study reports on surveys carried out in the austral springs of 2005 and 2006 using a simple and effective method for sampling from the sea-ice. These surveys enabled the evaluation of the spatial range of the nursery area of the Antarctic silverfish in the sea-ice of the coastal area of Victoria Land between the Coulman Island and the Drygalski Glacier Tongue. P. antarcticum eggs were concentrated in an area of Terra Nova Bay of about 270?km², encompassing two adjacent sites, Gerlache Inlet and Silverfish Bay. The present results add information on life cycle and hatching period of the Antarctic silverfish and confirm the importance of the Terra Nova Bay as a nursery area for this important species. Moreover, the survey points to the sea-ice cover and platelet ice as important environmental features of the nursery area.     

Molecular and comparative analyses of type IV antifreeze proteins (AFPIVs) from two Antarctic fishes, Pleuragramma antarcticum and Notothenia coriiceps

Antifreeze protein type IV (AFPIV) cDNAs and genomic DNAs from the Antarctic fishes Pleuragramma antarcticum (Pa) and Notothenia coriiceps (Nc) were cloned and sequenced, respectively. Each cDNA encoded 128 amino acids, with 94% similarity between the two and 83% similarity with AFPIV of the longhorn sculpin, Myoxocephalus octodecemspinosus. The genome structures of both genes consisted of four exons and three introns, and were highly conserved in terms of sequences and positions. In contrast, the third intron of PaAFPIV had additional nucleotides with inverted repeats at each end, which appeared to be a MITE-like transposable element. Comparative analysis revealed that fish AFPIVs were widely distributed across teleost fishes, well conserved in their intron positions, but more variable in intron sequences and sizes. However, the intron sequences of two Antarctic fishes were highly conserved, indicating recent radiation of notothenioids in the evolutionary lineage. The recombinant PaAFPIV and NcAFPIV were expressed in E. coli, and examined antifreeze activity. PaAFPIV and NcAFPIV gave ice crystals with star-shaped morphology, and thermal hysteresis (TH) values were 0.08°C at the concentration of 0.5mg/ml.     

Food and feeding ecology of postlarval and juvenile Pleuragramma antarcticum (Pisces; Notothenioidei) in the seasonal pack ice zone off the Antarctic peninsula

Summary Stomach contents of 275 postlarval and 269 one year old juvenile Pleuragramma antarcticum caught in February 1982, March 1981 and November 1977 in the Bransfield Strait and adjacent waters were investigated. Juveniles in November 1977 fed mainly on calanoid copepods and Oithona spp. The principal food of postlarvae in February 1982 were Oncaea spp., eggs of calanoid copepods and tintinnids whereas the staple food of juveniles in February 1982 consisted of calanoid copepods of which copepodites of Calanus propinquus were most abundant. Eggs of Euphausia superba were frequently ingested by postlarvae and were the main component of the juvenile diet in the Antarctic Sound. Postlarvae in March 1981 preyed on Oncaea spp. and calanoid eggs. There was a shift from feeding on cyclopoid copepods by postlarvae to feeding on calanoids by juveniles. The food particle size increased as a function of fish length. Maximum prey size was controlled by mouth width up to at least 45 mm standard length. Prey selection was apparently food density dependent with an inverse relationship between food abundance and selective feeding. Density dependent size selection may have resulted in resource partitioning among postlarval and juvenile fishes in February. This specific feeding behaviour is likely to be an adaptation to the low zooplankton stocks of the high Antarctic shelf waters to minimize food competition among the youngest age classes. Predation on krill eggs indicated that the frequent association of one year old juvenile Pleuragramma antarcticum with the Antarctic krill is related to local spawning events.     

Feeding plasticity of the Antarctic fish Trematomus hansoni Boulenger, 1902 (Pisces: Nototheniidae): the influence of fishery waste on the diet

The consumption of fishery waste by the Antarctic bottom-dwelling fish, Trematomus hansoni, is described in the Cosmonaut Sea. The study indicates that T. hansoni is able to switch its feeding habits at the time of fishery operations by incorporating fishery waste into the diet. Received: 22 September 1997 / Accepted: 21 November 1997     

Insights into spatial distribution patterns of early stages of the Antarctic silverfish,Pleuragramma antarctica,in the platelet ice of Terra Nova Bay,Antarctica

Polar Biology - The Antarctic silverfish, Pleuragramma antarctica, is ecologically important for connecting lower and upper trophic levels within the coastal marine Antarctic food web. In recent...     

Distribution and abundance of postlarval and juvenile Pleuragramma antarcticum (Pisces, Nototheniidae) off Terra Nova Bay (Ross Sea, Antarctica)

During the Italian Antarctic Expedition of 1987–1988 zooplankton was collected in Terra Nova Bay (Ross Sea) by a multinet BIONESS (250-μm or 500-μm net mesh size). The early life history of Pleuragramma antarcticum was described from 268 samples. More than 98% of a total of 34,436 fish larvae belonged to P. antarcticum. The mean relative abundance in the whole area for positive 0 to 150-m hauls was 434 ind./100 m³ (±720 SD). Postlarvae were most abundant and frequent in the samples (99.8%) while low concentrations of juveniles were found (n=67). Length of age group 0 ranged from 8 to 20 mm and age group 1+ were from 36 to 53 mm. Average growth rate over a period of 1 year was 0.08 mm per day. Based on modal values, the mean daily increment of P. antarcticum postlarvae in the period 5 January to 2 February (29 days) was about 0.21 mm. Highest abundances occurred near Cape Washington (mean: 2,108 ind./100 m³) while lower densities were recorded in the northern basin of the self (31 ind./100 m³). Horizontal and vertical distribution patterns in Terra Nova Bay seem to be strictly correlated to hydrographic features and different water masses with highest densities associated with the westward flowing current of the limb of the Antarctic coastal current and southern limb of the Ross Sea Gyre. These currents become part of the clockwise gyre in Terra Nova Bay. More than 62% of the postlarvae were collected in well-stratified warm surface water (0–50 m) near the summertime thermocline (20–70 m). The northern part of Terra Nova Bay seems to represent nursery ground of early stages of P. antarcticum and the presence of permanent polynya could provide favourable food conditions for development of the first stages of life. Differing distribution patterns probably reflect an interaction of various parameters including bathymetry, floating ice shelf, hydrographic features such as currents, local eddies and frontal systems, with P. antarcticum postlarval biology (spawning) and ecology (feeding, horizontal and vertical distribution patterns). Received: 10 June 1996 / Accepted: 19 July 1997     

Spatial distribution of Pleuragramma antarcticum (Pisces: Nototheniidae) near the Filchner- and Larsen ice shelves (Weddell sea/antarctica)

Summary In the zooplankton collected during three German Antarctic expeditions to the southern Weddell Sea in 1979/80, 1980/81 and 1983, the post-larvae of the nototheniid fish Pleuragramma antarcticum were found in 71–94% of the samples and represented 85–98% in numbers of all fish caught. In 1983, the abundance of post-larvae was up to 88 ind./m² (corresponding to 3 ind./m³). Highest concentrations were generally found over the continental slope and innershelf depressions. More than 70% of the post-larvae were caught in the upper 50–100 m water layer, in the well stratified Summer Water of -1.3 to -0.5°C. The Summer Water is shifted towards the outer edge of the shelf by Ekman transport and accumulation of post-larvae in the slope front and eddies can be explained by this drift. Older Pleuragramma antarcticum of age 1 resemble in their vertical distribution the juveniles and adults, 40–60% of which were caught on the shallower parts of the shelf in cold Ice Shelf Water of -1.8 to -2.1°C in depths below 200 m. Mean abundances of the yearclasses varied by a factor of 16 in age 0 fish and by factor of 10 in age 1. The size of a yearclass may be related to the varying appearance and persistence of an ice free coastal polynya.     

The presence and quantification of splenic ice in the McMurdo Sound Notothenioid fish, Pagothenia borchgrevinki (Boulenger, 1902)

Survival of some polar fishes is associated with high levels of circulating antifreeze glycoproteins (AFGPs). AFGP prevent ice growth giving rise to thermal hysteresis. The inhibiting action of AFGPs implies that polar fish contain ice to which AFGPs adsorb. Cryopelagic Pagothenia borchgrevinki, inhabiting the ice-laden waters of McMurdo Sound, Antarctica, were assayed for ice and ice was found on skin, gills, in the intestine, and in the spleen. Two methods used to assess the number of ice crystals in spleens gave comparable results (12.1 +/− 1.9 and 22 +/− 3.8 per spleen). Attempts were made to measure the rate of uptake of ice by P. borchgrevinki held in cages immediately beneath the sub-ice platelet layer in McMurdo Sound; uptake was sporadic. Introduction of ice into fish by spray freezing a small patch of the integument resulted in detection of splenic ice after 1 h, illustrating that a mechanism exists for ice transport from the periphery to the spleen. Splenic ice did not seem to be eliminated from fish held in ice-free water at − 1.6 °C for approximately two months. The relatively small number of splenic ice crystals and the slow rate of ice uptake suggest efficient ice barriers exist in P. borchgrevinki.     

Purification and biochemical characterization of a cadmium metallothionein from the digestive gland of the Antarctic scallop Adamussium colbecki (Smith, 1902)

A cadmium-binding protein was purified from the digestive gland of the Antarctic scallop, Adamussium colbecki, and biochemically characterized. Purification procedures included gel permeation and anion exchange chromatography, followed by preparative polyacrylamide gel electrophoresis. Our results demonstrate that the A. colbecki cadmium-binding protein has the general properties of metallothioneins: low molecular weight of about 10 kDa, spectroscopic features typical of cadmium thiolate clusters and high metal (cadmium) content. Analysis of amino acid composition reveals the absence of aromatic amino acids, histidine, methionine and arginine. Asparagine and glutamine are also absent. The A. colbecki metallothionein shows high levels of glycine (14%), aspartic acid (14%), glutamic acid (11%) and a low lysine content (4%); the A. colbecki metallothionein shows a lower cysteine content (12%) compared to other metallothioneins (17–30%) purified from both vertebrate and invertebrate organisms. The presence of a metallothionein in the digestive gland of A. colbecki suggests that in cold-ocean-adapted molluscs the heavy metal homeostasis mechanisms may have evolved similarly to those of organisms living in temperate marine environments, although the A. colbecki cadmium-binding protein shows a typical amino acidic composition that might reflect a peculiar physiological role. Accepted: 7 August 2000     

Some Antarctic and sub-Antarctic species of Celleporidae (Bryozoa, Cheilostomata)

Sixteen species of cheilostome Bryozoa, within the family Celleporidae Busk, 1852, are reported from Antarctica and the sub-Antarctic south-west Atlantic. Nine new species are described in the genera Osthimosia Jullien, 1888, Celleporina Gray, 1848 and Turbicellepora Ryland, 1963. Spigaleos gen. nov. is introduced for Cellepora horneroides Waters, 1904.