Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

Proximate composition of Antarctic mesopelagic fishes

Eleven mesopelagic fish species from the Weddell/Scotia Sea region of the Antarctic captured during the austral spring 1983, austral fall 1986, and austral winter 1988, were analyzed for proximate composition. Water, ash level, protein, lipid and carbohydrate were examined in relation to depth of occurrence and season. No depth-related trends were evident, primarily due to a low species diversity and minimal differences in those species' vertical distributions. The Antarctic speciesElectrona antarctica showed a significant increase in lipid level (% wet wt and % ash-free dry wt) between spring, fall and winter. The increase may signify an accumulation over the productive season, possibly as a reserve for the winter months. Lipid levels (% wet wt and % ash-free dry wt) were significantly lower in the Weddell Sea specimens examined in this study than in previously examined identical and congeneric species taken during the same season from a more productive near-shore Antarctic region. Comparisons with congeners and confamilials from tropical-subtropical and temperate systems revealed variable trends. The Antarctic speciesE. antarctica andCyclothone microdon had lower water and protein (% wet wt) levels than similar species from tropical-subtropical or temperate regions. Lipid levels of the two species are similar to temperate individuals, while energy levels are slightly higher. In contrast, species of the genusBathylagus show no trends in composition as a function of latitude. Differences in productivity, water-column temperature-structure, and seasonality are important considerations when examining trends among mesopelagic species.     

Age and growth of Electrona antarctica (Pisces: Myctophidae), the dominant mesopelagic fish of the Southern Ocean

Numerically and in biomass, the lanternfish Electrona antarctica is the dominant fish in the vast pelagic region of the Southern Ocean bounded on the north by the Antarctic Convergence and in the south by the Antarctic continental shelf. It is an important krill predator, and in turn is important in the diets of flighted and swimming seabirds. Further, it is the southernmost and coldest-dwelling representative of the globally distributed fish family Myctophidae. The present study was undertaken to estimate the species' growth rate and average life span, to incorporate the information in a basic energy budget, and to compare the growth of E. antarctica with more northerly confamilials. Fishes were aged using primary growth increments that were resolved on sagittal otoliths using three sequential techniques: thin-section grinding and polishing, etching, and scanning electron microscopy (SEM). Based on increment width (0.8 to 1.2 μm), continuity, and previous studies on confamilials, the microincrements were assumed to be deposited on a daily basis. Montages of SEM photomicrographs were constructed for each sagitta to allow the daily rings to be counted over the entire life span of 31 individuals representing the entire size range of the species. Results suggest a larval stage of 30 to 47 d and a maximum life span of 3.5 yr, with females growing faster than males in the last 1.5 yr of life and reaching a larger maximum size. Construction of a simple energy budget using the best information available suggests that a surplus of energy is available to support the observed growth rates (0.05 to 0.07 mm d⁻¹). The results of the present study contrast markedly with previous estimates of an 8 to 11 yr maximum age for E. antarctica. These results provide important data addressing the ecology and population dynamics of the pelagic Antarctic ecosystem. E. antarctica is the end-member species in the continuum of vertically migrating myctophids that extend from the equator to the polar circle. Its growth rate is consonant with that of all other myctophid species examined using primary growth increments to determine age. The present study, in conjunction with earlier studies, suggests that growth rates of mesopelagic species are far higher than previously thought. Received: 12 September 1997 / Accepted: 25 July 1998     

Lipid composition of twenty-two species of Antarctic midwater zooplankton and fish

Total lipids, hydrocarbons, wax esters, triacylglycerols, and phospholipids were determined for 22 major biomass species of zooplankton and fish in an Antarctic mesopelagic community that were collected in 1982 and 1983. Lipid levels were similar to levels in more temperate mesopelagic species. Total lipid concentration was depth related, with all lipid-rich species being collected at depths greater than 230 m. Wax ester content in copepods (60 to 70% of the total lipid) was generally higher than in subtropical species. Lipids indicated three predator-prey relationships (Parandania boecki-Atolla wyvillei, Thysanoessa macrura-Calanoides acutus andEurythenes gryllus-Atolla wyvillei). These were confirmed by gut content analyses. The mesopelagic fishBathylagus antarcticus, Pleuragramma antarcticum, andProtomyctophum bolini stored most lipid intramuscularly, whereasElectrona antarctica andGymnoscopelus nicholsi contained extensive stores in subcutaneous lipid sacs. The intramuscular lipids inP. antarcticum and the subcutaneous lipid sacs ofE. antarctica were primarily wax esters, possibly used for increased buoyancy or long-term energy storage. Unlike the odd-carbon preference of aliphatic hydrocarbons which typifies terrestrial plants and temperate marine organisms, even-carbon chain-length paraffins predominate in 80% of the Antarctic species analyzed. Although the source of these even-carbon n-alkanes cannot be determined from our data, their dominance in the species analyzed suggests that an unusual biochemical pathway may be responsible for their synthesis in this ecosystem.     

Population structure of two Antarctic ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, in winter sea ice

The spatial distribution and population structure of two dominant ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, were studied during winter at nine locations in east Antarctic fast ice. These species accounted for at least 90% of the total metazoan abundance at each location. Abundances were high, reaching 175 individuals l^-1 (190,000 m^-2) for D. glacialis and 660 l^-1 (901,000 m^-2) for P. antarctica. These abundances were probably partly supported by the high biomass of ice-algae (Pearson correlation coefficient, r=0.75), as indicated by chlorophyll-a concentrations (1.7-10.1 µg l^-1). The population structures of each species suggested very different life-history strategies. All developmental stages of D. glacialis were isolated from the ice cores, including females with egg sacs, supporting the hypothesis that this species reproduces in the sea ice during winter. This strategy might assist D. glacialis in leading a continually colonising existence, whereby it responds opportunistically to the availability of favourable habitat patches. The populations of P. antarctica were composed primarily of nauplii (>99%), consistent with past observations of a synchronised life cycle for this species. The strong coupling of the developmental cycle of P. antarctica to the growth and decay of sea ice suggests that local extinctions might occur in areas where ice break-out is unpredictable.     

Ultrastructure of pedal muscle as a function of temperature in nacellid limpets

Temperature and mitochondrial plasticity are well studied in fishes, but little is known about this relationship in invertebrates. The effects of habitat temperature on mitochondrial ultrastructure were examined in three con-familial limpets from the Antarctic (Nacella concinna), New Zealand (Cellana ornata), and Singapore (Cellana radiata). The effects of seasonal changes in temperature were also examined in winter and summer C. ornata. Stereological methods showed that limpet pedal myocytes were 1–2 orders of magnitude smaller in diameter (≈3.5 μm) than in vertebrates, and that the diameter did not vary as a function of temperature. Mitochondrial volume density (Vv_(mt,f)) was approximately 2–4 times higher in N. concinna (0.024) than in the other species (0.01 and 0.006), which were not significantly different from each other. Mitochondrial cristae surface density (Sv_(im,mt)) was significantly lower in summer C. ornata (24.1 ± 0.50 μm² μm⁻³) than both winter C. ornata (32.3 ± 0.95 μm² μm⁻³) and N. concinna (34.3 ± 4.43 μm² μm⁻³). The surface area of mitochondrial cristae per unit fibre volume was significantly higher in N. concinna, due largely to the greater mitochondrial volume density. These results and previous studies indicate that mitochondrial proliferation in the cold is a common, but not universal response by different species from different thermal habitats. Seasonal temperature decreases on the other hand, leading preferentially to an increase in cristae surface density. Stereological measures also showed that energetic reserves, i.e. lipid droplets and glycogen in the pedal muscle changed greatly with season and species. This was most likely related to gametogenesis and spawning.     

Comparative ecology of Lessonia nigrescens and Durvillaea antarctica (phaeophyta) in Central Chile

The Lessonia nigrescens-Durvillaea antarctica belt is the most conspicuous feature of the intertidal-subtidal boundary in Central Chile, with L. nigrescens attaining larger cover and biomass values than D. antarctica. Human predation of the otherwise competitively dominant D. antarctica has been suggested as accounting for its scarcity. Testing of this hypothesis included field studies of species distribution in places with and without human accessibility and various degrees of wave impact, comparative morphometric and population studies, evaluation of the ecologic role of the limpet-kelp association and comparative measurements of growth, reproduction and survival capacity. Results indicate that L. nigrescens and D. antarctica have morphologies typically adapted to habitats with different wave shock. Contrary to previous predictions, L. nigrescens appears as a plant better adapted for places with strong wave impact, which are the most common in Central Chile. Complete space monopolization by L. nigrescens is prevented by a series of adaptations of D. antarctica. Certain morphological forms of this species are less affected by wave action, allowing a population stock to persist even at the most exposed places. Boring into algal holdfasts by invertebrates weakens the mechanical resistance of old, eroded plants providing open space where juveniles of either species could settle. D. antarctica seems to take greater advantage of this primary space by means of a fugitive life history. Finally, the survival of D. antarctica increases if attached to the top of L. nigrescens holdfasts. It is therefore concluded that result of the interaction between these 2 brown algae depends on the degree of water impact. In sheltered habitats competitive displacement of L. nigrescens could occur but it is unlikely to be a continuous process. In exposed habitats D. antarctica is constantly removed by water movement, but persists because of a higher colonization rate.     

Genetic diversity of Nymphon (Arthropoda: Pycnogonida: Nymphonidae) along the Antarctic Peninsula with a focus on Nymphon australe Hodgson 1902

Sea spiders are conspicuous, and often abundant, members of the Antarctic benthic community. Nymphonidae (Pycnogonida) in Southern Ocean waters comprise over 240 species which are often difficult to assign due to their intraspecific ‘highly variable’ morphology. In particular, Nymphon australe, the numerically dominant species in Antarctic waters is known to have a high level of phenotypic variation in external morphology and is also reported to have a circumpolar distribution. Circumpolarity seems contradictory to the pycnogonid’s brooding lifestyle and presumably limited dispersal. Here we examine the genetic diversity of several Nymphon species collected in the Antarctic Peninsular region. Concomitantly, we assess the genetic structure of N. australe to gain insight into Nymphon dispersal capacity. Cytochrome c oxidase subunit I (COI) and 16S ribosomal gene data suggest a recent common history and/or recent gene-flow of N. australe populations across nearly 800 km of the Antarctic Peninsula. Furthermore, these data support that the Antarctic Peninsula region may hold two previously unrecognized species of Nymphon. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

On the lipid biochemistry of polar copepods: compositional differences in the Antarctic calanoids Euchaeta antarctica and Euchirella rostromagna

During austral summer of 1985 different developmental stages (CIII, CIV, CV, females, males) of the Antarctic copepod Euchaeta antarctica and females of Euchirella rostromagna were collected in the southeastern Weddell Sea to determine their lipid contents and compositions. For E. antarctica the analyses revealed a strong ontogenetic accumulation of lipids towards the older copepodids with highest lipid contents in late CV stages and adults. The females of E. rostromagna had moderate lipid levels. The most striking difference between these two species concerns their lipid class compositions. E. antarctica deposited predominantly wax esters, whereas in E. rostromagna the major lipid class consisted of triacylglycerols, an unusual storage lipid in polar marine copepods. Principal fatty acids in E. antarctica were the monounsaturates 18:1(n-9) and 16:1(n-7), especially in the lipid-rich stages, while the polyunsaturated fatty acids 20:5(n-3) and 22:6(n-3), usually membrane lipids, dominated in the lipid-poor stages. The wax ester moieties in E. antarctica consisted almost entirely of 14:0 and 16:0 fatty alcohols. Major components in E. rostromagna were the fatty acids 18:1(n-9), 16:0, 20:5(n-3) and 22:6(n-3). The potential of fatty acids and alcohols as typical trophic markers is rendered largely insignificant in the two species due to catabolic processes.     

Changes in amphipod densities among macroalgal habitats in day versus night collections along the Western Antarctic Peninsula

Amphipods along the western Antarctic Peninsula appear to gain refuge from predators by associating with chemically defended macroalgae rather than palatable macroalgae. However, nothing is known about amphipod activity at night. If foraging on non-chemically defended macroalgae regularly occurs, then nocturnal foraging seems beneficial since visual predators are disadvantaged. To test this hypothesis, we collected three common macroalgal species and affiliated mesograzers, approximately 3 h before and after sunset. All associated mesofauna were counted and densities recorded. Amphipod densities were significantly decreased during the night on the chemically defended Desmarestia menziesii, while significantly increased on the palatable Iridaea cordata. Additionally, the amphipod Gondogeneia antarctica was found in significantly higher densities at night on Palmaria decipiens, a species shown to be readily eaten by G. antarctica and omnivorous fish. We believe that chemically defended macroalgae act as a refuge for mesograzers during the day, while more widespread foraging occurs at night.     

Feeding in dominant Antarctic copepods—does the morphology of the mandibular gnathobases relate to diet?

This study should clarify the importance of morphology and stability of the mandibular gnathobases for the diet of Antarctic copepod species. The gnathobase morphology of the dominant copepod species Calanoides acutus, Calanus propinquus, Ctenocalanus citer, Rhincalanus gigas, Metridia gerlachei, Stephos longipes, Microcalanus pygmaeus and Paraeuchaeta antarctica from the Southern Ocean was investigated in detail by means of a scanning electron microscope. The mandibular gnathobases of C. acutus, C. propinquus and C. citer have relatively short and compact teeth. These species feed mainly on diatoms and are able to crack the silicious diatom frustules with their mandibular gnathobases by directed pressure. In contrast the teeth of the mandibular gnathobases of P. antarctica are very long and pointed. The nutrition of this species consists predominantly of other smaller copepod species. The motile prey can be held by skewering, using the gnathobases, and then eventually minced. The mandibular gnathobases of P. antarctica have notably more small bristles than those of the other investigated copepod species. These bristles are probably associated with receptors and could serve to locate the prey. The morphology of the gnathobases of R. gigas and M. gerlachei is between that of P. antarctica on the one side and that of C. acutus, C. propinquus and C. citer on the other. Based on the morphology of its gnathobases the copepod species S. longipes, which has to date been found to feed primarily on phytoplankton, mainly ice algae, must also be considered a zooplankton feeder. The investigation showed that M. pygmaeus has gnathobases with surprisingly long and pointed teeth, indicating that this species very probably feeds both on phyto- and on zooplankton organisms. While the mandibular gnathobases of the males of C. propinquus, R. gigas, M. gerlachei and S. longipes have the same morphology as the females of the respective species, in the other four investigated copepod species the males have reduced (C. acutus, C. citer and M. pygmaeus) or completely missing mandibular gnathobases (P. antarctica). The teeth of the gnathobases of all studied species with the exception of M. gerlachei consist of a different material than the remaining parts of the gnathobases. This material seems to be silicate, which probably enhances the stability of the gnathobase teeth.Communicated by O. Kinne, Oldendorf/Luhe     

Macrobenthic species assemblages in Ellis Fjord,Vestfold Hills,Antarctica

A study was made of the sub-littoral benthic environment of Ellis Fjord, a 10 km-long fjord located near Davis Station, in the Vestfold Hills, Antarctica, over a 15 mo period (November 1984 to February 1986). Data were collected by SCUBA diving and underwater photography and were inhitially analysed by ordination techniques (non-metric multidimensional scaling). Ordinations showed substratum type to be the factor most strongly associated with changes in the distribution and abundance of macrobenthic species within the fjord. Other factors shown to be associated with changes in macrobenthic species assemblages were depth, distance from the fjord mouth, bottom slope, shoreline characteristics, current speed, and the presence of low-salinity water at shallow depths during the summer melt. The four major substratum types in Ellis Fjord were sand, rock, Serpula narconensis colonies and Phyllophora antarctica thalli. S. narconensis colonies supported the most species and sand substrate supported the least. P. antarctica is the only macrophyte species which occurred in the fjord. S. narconensis colonies in Ellis Fjord from one of the largest known tubeworm reefs in the world. The assemblages of benthic species in Ellis Fjord were different from those seen at other sub-littoral benthic sites off the Vestfold Hills, and at other Antarctic sites. There was a far greater proportion of filter-feeding species in the fjord than at other sub-littoral benthic sites off the Vestfold Hills. Factors which are thought to have caused these differences are the high level of organic but low level of inorganic input into the benthic system of the fjord, and the absence of anchor ice from the fjord.     

Genetic differentiation between morphotypes in the Antarctic limpet <Emphasis Type="Italic">Nacella concinna</Emphasis> as revealed by inter-simple sequence repeat markers

The limpet Nacella concinna (Strebel 1908) was the focus of numerous studies dealing with Antarctic benthos. One of the main characteristics of the species is the presence of two distinguishable morphotypes, one inhabiting the intertidal (during summer) and the other inhabiting the subtidal. For a long time these forms were considered as an expression of phenotypic plasticity, since previous studies did not found genetic differences between them. In the present work, we performed both a morphometric and a genetic differentiation analysis (using ISSR-PCR markers) of these two forms in three stations sampled at Potter Cove, South Shetland Islands. The results confirmed the morphological differences between intertidal and subtidal forms reported in other Antarctic localities. The genetic differences detected indicate that the two forms can be considered as genetically distinct populations maintaining low levels of gene flow. The degree of reproductive isolation of the ecotypes is discussed, as well as the possible origin of the divergence. The genetic differentiation observed can also have behavioral and physiological correlates, pointing out the importance of taking into account the potential differences in the response of both populations to different conditions in future studies in this species. M. C. de Aranzamendi and R. Sahade contributed equally to this paper.     

Krill evolution and the Antarctic ocean currents: evidence of vicariant speciation as inferred by molecular data

The phylogenetic relationships of the Antarctic krill Euphausia superba, the key species in the Antarctic food web, and other Antarctic and sub-Antarctic cuphausiids have been investigated using the 16S ribosomal mitochondrial gene. The phylogenetic reconstructions indicated that the Antarctic species form a monophyletic clade separated by the non-Antarctic species. The results revealed a large genetic divergence between the Antarctic (E. superba and E. crystallorophias) and sub-Antarctic species (E. vallentini). The time of separation between these species, estimated from the molecular data, is around 20 million years ago, which is comparable with the geological time of the formation of a circum-Antarctic water circulation and the Antarctic Polar Frontal Zone. The euphausiid molecular phylogeny therefore represents evidence for vicariant speciation.     

No evidence for genetic differentiation between Antarctic limpet Nacella concinna morphotypes

The extent to which genetic divergence can occur in the absence of physical barriers to gene flow is currently one of the most controversial topics in evolutionary biology, with implications for our understanding of speciation, phenotypic plasticity and adaptive potential. This is illustrated by a recent study reporting a surprising pattern of genetic differentiation between intertidal and subtidal morphotypes of the broadcast-spawning Antarctic limpet Nacella concinna. To explore this further, we collected almost 400 Antarctic limpets from four depths (intertidal, 6, 15 and 25 m) at Adelaide island, Antarctica, and conducted a combined morphometric and genetic analysis using 168 polymorphic amplified fragment length polymorphism (AFLP) loci. Morphological analysis revealed not only pronounced differences between the two morphotypes, but also a continuous cline in shell shape from the intertidal zone down to 25 m depth, suggesting that the distinction between the morphotypes may be artificial. Moreover, genetic analysis using both F _st and a Bayesian analogue found no evidence for differentiation either between the two morphotypes or by depth, and a Bayesian cluster analysis did not detect any cryptic genetic structure. Our findings lend support to the notion that limpets can be phenotypically highly plastic, although further studies are required to determine unequivocally whether there is any genetic basis to the observed variation in shell morphology.     

Fatty acid signature analysis documents the diet of five myctophid fish from the Southern Ocean

Fatty acid (FA) and fatty alcohol (FAlc) compositions of both total lipid and neutral lipid fractions were studied for five myctophid species sampled in Kerguelen waters. Both qualitative and quantitative FA signature analyses were then performed to investigate their diet over longer time scales than the conventional stomach content analysis. Regarding their lipid class, FA and FAlc compositions, the five species could be discriminated into two groups: wax-ester-rich species (Electrona antarctica, Krefftichthys anderssoni) characterised by large amounts of monounsaturated FAs (>73% of total FAs) and triacylglycerol-rich species (Electrona carlsbergi, Gymnoscopelus nicholsi, Protomyctophum bolini) with major amounts of saturated and monounsaturated FAs (>29 and >46% of total FAs, respectively). Qualitative and quantitative FA analyses showed that K. anderssoni mainly preyed upon copepods, E. antarctica upon copepods and more euphausiids and P. bolini and E. carlsbergi mainly upon euphausiids with some copepods, while G. nicholsi had a more diverse diet. This study shows the usefulness of quantitative statistical analysis to determine the diet of Antarctic and sub-Antarctic predators and stresses the need of increasing the lipid and FA analyses of more zooplanktonic and micro-nektonic marine species.     

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus in blue (Mytilus galloprovincialis) and greenshell (Perna canaliculus) mussel populations along a salinity gradient

In several marine bivalve species, biochemical genetic variation at a leucine aminopeptidase (LAP) locus is associated with environmental variability, primarily salinity fluctuation. Population genetic variation at a LAP locus was investigated here in two sympatric mussel species (Mytilus galloprovincialis and Perna canaliculus) from three locations along a salinity gradient in Wellington Harbour, New Zealand. The data for M. galloprovincialis and P. canaliculus do not support the hypothesis that the LAP polymorphism in either species is associated with salinity variation among adult mussels. Due to the absence of small mussels among the samples it is not possible to discount the hypothesis that selection acts primarily against juveniles, as it does for M. edulis in Long Island Sound, USA. Wellington Harbour populations of M. galloprovincialis exhibited large and often highly significant heterozygote excesses at the LAP locus, whereas populations of P. canaliculus from the same locations exhibited large and highly significant heterozygote deficiencies. The reason for this inter-specific difference in population structure is unknown. If it is the result of selection, this suggests that selection acts differentially upon the two species, because demographic attributes and reproductive biology are very similar in the two species. For both M. galloprovincialis and P. canaliculus, significant levels of population genetic heterogeneity were recorded among three locations separated by only 8 to 12 km. Neither species exhibited shell length-dependent genetic variation at the LAP locus, suggesting that for these two species the LAP polymorphism is not associated with variation in shell length. Received: 30 December 1996 / Accepted: 6 January 1997     

Longitudinal and vertical trends in stable isotope signatures (δ13C and δ15N) of omnivorous and carnivorous copepods across the South Atlantic Ocean

Stable isotope (SI) ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were measured in omnivorous and carnivorous deep-sea copepods of the families Euchaetidae and Aetideidae across the Atlantic sector of the Southern Ocean to establish their trophic positions. Due to high and variable C/N ratios related to differences in lipid content, δ¹³C was corrected using a lipid-normalisation model. δ¹⁵N signals ranged from 3.0–6.9‰ in mesopelagic species to 7.0–9.5‰ in bathypelagic congeners. Among the carnivorous Paraeuchaeta species, the epi- to mesopelagic species Paraeuchaeta antarctica had lower δ¹⁵N values than the mesopelagic P. rasa and bathypelagic P. barbata. The same trend was observed among omnivorous Aetideidae, but was not significant. In the most abundant species P. antarctica, individuals from the western Atlantic had higher δ¹³C and δ¹⁵N values than specimens at the eastern stations. These longitudinal changes in δ¹³C and δ¹⁵N values were attributed to regional differences in hydrography and sea surface temperature (SST), in particular related to a northward extension of the Antarctic Polar Front (APF) at the easternmost stations. The results indicate that even in a mesopelagic carnivorous species, the changes in surface stable isotope signatures are pronounced.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

Spatio-temporal changes in diversity and community structure of planktonic copepods in Sagami Bay, Japan

Seasonal changes in diversity and community structure of planktonic copepods at a shelf site in Sagami Bay, Japan was studied in relation to cross-shelf interaction of species components. Seasonal mesozooplankton samples were collected from the shelf station (St. M) of the north-west part of Sagami Bay from 1995 to 1997. Vertical multi-layered samples were collected near the center of Sagami Bay (St. P) in June 1996. A total 185 copepod species were identified from the two stations. We observed a clear seasonal succession in calanoid diversity and community structure at St. M from a simple shelf water community (>11 species) during spring blooming periods to highly diverse and mixed communities (ca 20–30 species) of shelf water species coupled with various Kuroshio Current species during late summer to autumn. Cluster and non-metric multidimensional scaling ordination analyses showed two distinct calanoid community groups. One group, which included samples of St. M and the surface layer of St. P, consisted of shelf water species, such as Calanus sinicus, Ctenocalanus vanus, Paracalanus spp., and Kuroshio species, such as, Canthocalanus pauper, Scolecithrix danae, etc. The other cluster was restricted to the samples collected from mid and deep layers at St. P, which consisted of meso- and bathypelagic species and Oyashio species (cold-current species, such as Neocalanus cristatus, Pseudocalanus spp., Eucalanus bungii and Metridia pacifica). In the mid and deep layers at St. P, the population of dormant copepodid stage V (CV) of Eucalanus californicus and C. sinicus were dominant. The deep CV population of C. sinicus might be ecologically discriminated from the surface and shelf water population due to their larger body length and dormant life cycle. E. californicus was also collected at the shelf site during each spring bloom period, whereas the population might descend into the mid- and deep-layers of the central bay before summer. Our results suggest that the seasonal fluctuation of community structure in the shelf water was controlled by both physical (Kuroshio Current) and biological factors, i.e., spring bloom and ontogenetic vertical migration of E. californicus. In particular, transport and diffusion processes of Kuroshio Current in Sagami Bay played a key role in controlling the shelf water calanoid community.