The diets of polar cod (Boreogadus saida) from August 2008 in the US Beaufort Sea

Polar cod (Boreogadus saida) play an integral part in the Arctic ecosystems linking the upper and lower trophic levels. Though their estimated biomass is considerable, recent knowledge of their diets in the US Beaufort Sea is sparse. Collections of polar cod from the US Beaufort Sea were made during August 2008 using demersal and pelagic trawls. Polar cod diet composition was quantified as percent prey weight, percent prey count, and frequency of occurrence of prey. The diet composition between the demersal- and pelagic-captured cod showed differences in all these categories. Polar cod captured in the demersal nets primarily fed on fish (by weight), and pelagic cod primarily fed on copepods (frequency of occurrence) and euphausiids (by weight). In general, these dominant preys are different than what has been reported in other studies describing polar cod diets.     

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.     

Characteristics of under-ice swarming of polar cod Boreogadus saida (Gadidae) in the Central Arctic Ocean

Phenological observations on under-ice swarming behavior of Polar cod Boreogadus saida were performed during the winter period 2009–2010 in the Canadian sector of the Arctic Ocean on the North Pole NP-37 drifting station. These observations and ichthyologic collections were the basis of the present study. The schools were mostly formed by the immature specimens (TL = 75–169 mm; 1+ to 4+ age), the dominating group was presented by fish of 2+ age (86–94% of total abundance). The driving factors of such large under-ice swarming of Polar cod B. saida in the Arctic are discussed, particularly, the interactions of Polar cod development and the ice drift in the studied region. This species inhabits the overcooled water (?1.8°C) under the ice, so the scheme of the antifreeze agent producing in the fish blood is considered. We make assumptions about under ice migrations of Polar cod swarming.     

Functional characterization of parvalbumin from the Arctic cod (Boreogadus saida): similarity in calcium affinity among parvalbumins from polar teleosts

Calcium dissociation constants (KD) were measured as a function of temperature for parvalbumin, a small acidic protein expressed abundantly in fast-twitch muscle, from the Arctic cod (Boreogadus saida) and compared to values previously determined for Antarctic and temperate zone teleosts. Estimates of KD were derived independently from fluorometric titrations and calorimetry. In addition, the primary structure of B. saida parvalbumin was determined. Calcium KDs for parvalbumin from B. saida were fundamentally similar to those for parvalbumins from Antarctic species (6.68+/-0.59 nM and 7.77+/-0.72 nM at 5 degrees C, respectively), but significantly different from temperate zone species (1.35+/-0.28 nM at 5 degrees C). However, estimates of KD for B. saida parvalbumin at 5 degrees C closely matched values for temperate zone fish at 25 degrees C (6.54+/-0.56 nM), recapitulating the prior observation that calcium affinity of parvalbumin is conserved at the native temperature of teleost fish. Full sequence of B. saida parvalbumin was generated using reverse-phase HPLC and RACE-PCR. The Arctic parvalbumin showed 83% homology to a carp parvalbumin. None of the 16 total substitutions between the two parvalbumins resided in the cation binding sites of the protein, indicating that the structural locus of the thermal sensitivity of function lies outside the active regions.     

Sub-lethal levels of waterborne petroleum may depress routine metabolism in polar cod Boreogadus saida (Lepechin, 1774)

Petroleum-related activities in Arctic waters are rapidly increasing parallel to the ongoing thinning of the Arctic sea ice. As part of a series of studies on petroleum-induced stress in polar cod Boreogadus saida, we tested the effects of acute (~60 min) and chronic (4 weeks) exposure to the water soluble fraction (WSF) of petroleum on whole body metabolism inferred from measurements of oxygen consumption rates. The exposure of polar cod to WSF leads to a statistically significant depression in routine metabolism in the order Control (0.260 mg O₂ g fish⁻¹ h⁻¹; N = 6) > Chronic (0.191 mg O₂ g fish⁻¹ h⁻¹; N = 6) > Acute (0.110 mg O₂ g fish⁻¹ h⁻¹; N = 2), decoupling of routine metabolism and body mass but possibly also to a partial metabolic compensation after 4 weeks of exposure. The results are reviewed in context with similar studies on Antarctic and non-polar fishes.     

Sourcing fatty acids to juvenile polar cod (Boreogadus saida) in the Beaufort Sea using compound-specific stable carbon isotope analyses

Juvenile polar cod (Boreogadus saida) are often found in close association with sea ice and represent an important trophic link in the Arctic food web. However, the proportional contribution of sea ice algal production via the sympagic food web to the diet of polar cod is unknown. To estimate the proportional contribution of fatty acids (FAs) from sea ice-derived particulate organic matter (i-POM) to the diet of juvenile polar cod, we used FA profiling and compound-specific stable carbon isotope analysis of individual FAs from juvenile polar cod collected from three regions in the Beaufort Sea. The δ¹³C values of the FAs 14:0, 16:4n-1, 18:0, 20:5n-3 and 22:6n-3 in the polar cod were found to most strongly resemble pelagic POM rather than i-POM. Results from isotope-mixing models using diatom FA markers indicated that the proportional contribution of FAs from i-POM to juvenile polar cod was ≤2 %, which suggests that juvenile polar cod had not sourced their FAs from i-POM. Thus, changes in sea ice coverage due to environmental change may not affect juvenile polar cod in regard to nutrients such as FAs but may still affect their populations by reducing critical shelter from predators.     

The aggregation of polar cod (Boreogadus saida) in the deep Atlantic layer of ice-covered Amundsen Gulf (Beaufort Sea) in winter

During the Circumpolar Flaw Lead System Study (CFL, 2007–2008), large aggregations of polar cod were detected in winter in the Amundsen Gulf (Western Canadian Arctic) using the EK60 echosounder of the CCGS Amundsen research icebreaker. Biomass estimated over 10 months reached a maximum of 0.732 kg m⁻² in February. Aggregations were encountered only in the presence of an ice cover from December to April. The vertical extent of the aggregations was dictated by temperature and zooplankton prey distribution. In winter, polar cod generally occupied the relatively warm deep Atlantic Layer (>0°C), but a fraction of the densest aggregations occasionally followed zooplankton prey up into the cold Pacific Halocline (−1.6 to 0°C). The diel vertical migration of polar cod was precisely synchronized with the seasonally increasing photoperiod. Throughout winter, polar cod aggregations migrated to progressively deeper regions (from 220 to 550 m bottom depths) in response to increasing light intensity, presumably to avoid predation by visual predators such as the ringed seal. Comparing Amundsen Gulf and Franklin Bay indicates that the entrapment of polar cod in embayments during winter is an important mechanism to provide marine mammal predators with dense concentrations of their main prey within their diving range.     

In-situ observations on the distribution and behavior of amphipods and Arctic cod (Boreogadus saida) under the sea ice of the High Arctic Canada Basin

The occurrence and behavior of sympagic amphipods and Arctic cod (Boreogadus saida) were studied in the High Arctic Canada Basin by diving under the ice at seven stations in summer 2002. Still images of video-transects were used to obtain animal abundances and information on the structure of the ice environment. Mean amphipod abundances for the stations varied between 1 and 23 individuals m^–2, with an increase towards the western part of the basin. The standard deviation within the 31–51 images analyzed per station was small (<1 individual m^–2). Gammarus wilkitzkii was found in low abundances, often hiding in small ice gaps. Small amphipods (Onisimus spp., Apherusa glacialis, and juveniles of all species) tended to move freely along the bottom of the floes. B. saida occurred in narrow wedges of seawater along the edges of melting ice floes at three stations in water depths of 10–50 cm and was never found under the ice. The fish occurred in schools of 1–28 per wedge. Fish were inactive and did not escape the approaching diver. Resting in the wedges may be a strategy to reduce energetic requirements and avoid predators.We dedicate this publication to Professor Dr. J. Lenz (Kiel University).     

Size,age and diet of polar cod,Boreogadus saida (Lepechin 1773), in ice covered waters

Summary Polar cod (Boreogadus saida) associated with drifting sea-ice were collected in the western Barents sea and north of Svalbard with dip-nets while SCUBA-diving in 1986 and 1987. Length-frequency measurements and otolith-readings suggested that the specimens were either one or two years old. The diet of fish from the western Barents sea (first-year ice) consisted mainly of copepods (Calanus finmarchicus, Calanus glacialis) and the hyperiid amphipod Parathemisto libellula. Fish collected north of the Svalbard archipelago (multi-year ice) had a more diverse diet, in which P. libellula and the sympagic amphipod Apherusa glacialis contributed more to the total diet biomass than copepods.     

Patterns of population subdivision, gene flow and genetic variability in the African wild dog (Lycaon pictus)

African wild dogs are large, highly mobile carnivores that are known to disperse over considerable distances and are rare throughout much of their geographical range. Consequently, genetic variation within and differentiation between geographically separated populations is predicted to be minimal. We determined the genetic diversity of mitochondrial DNA (mtDNA) control region sequences and microsatellite loci in seven populations of African wild dogs. Analysis of mtDNA nucleotide diversity suggests that, historically, wild dog populations have been small relative to other large carnivores. However, population declines due to recent habitat loss have not caused a dramatic reduction in genetic diversity. We found one historical and eight recent mtDNA genotypes in 280 individuals that defined two highly divergent clades. In contrast to a previous, more limited, mtDNA analysis, sequences from these clades are not geographically restricted to eastern or southern African populations. Rather, we found a large admixture zone spanning populations from Botswana, Zimbabwe and south-eastern Tanzania. Mitochondrial and microsatellite differentiation between populations was significant and unique mtDNA genotypes and alleles characterized the populations. However, gene flow estimates (Nm) based on microsatellite data were generally greater than one migrant per generation. In contrast, gene flow estimates based on the mtDNA control region were lower than expected given differences in the mode of inheritance of mitochondrial and nuclear markers which suggests a male bias in long-distance dispersal.     

Nuclear and chloroplast microsatellites reveal extreme population differentiation and limited gene flow in the Aegean endemic Brassica cretica (Brassicaceae)

Nuclear and chloroplast microsatellite markers were used to study population structure and gene flow among seven Cretan populations of the Aegean endemic plant species Brassica cretica (Brassicaceae). Both nuclear and chloroplast markers revealed exceptionally high levels of population differentiation (overall F(ST)=0.628 and 1.000, respectively) and relatively little within-population diversity (overall H(S)=0.211 and 0.000, respectively). Maximum-likelihood estimates of directional migration rates were low among all pairs of populations (average Nm=0.286). There was no evidence that differences in flower colour between populations had any influence on historical levels of gene flow. In addition, a haplotype network showed that all five chloroplast haplotypes found in the sample were closely related. Together, these results suggest that current patterns of diversification in B. cretica are mainly a result of genetic drift during the last half million years. The main conclusions from the present study are consistent with the prevailing hypothesis that plant diversification in the Aegean region is driven by random rather than adaptive differentiation among isolated populations.     

Habitat barriers limit gene flow and illuminate historical events in a wide-ranging carnivore, the American puma

We examined the effects of habitat discontinuities on gene flow among puma (Puma concolor) populations across the southwestern USA. Using 16 microsatellite loci, we genotyped 540 pumas sampled throughout the states of Utah, Colorado, Arizona, and New Mexico, where a high degree of habitat heterogeneity provides for a wide range of connective habitat configurations between subpopulations. We investigated genetic structuring using complementary individual- and population-based analyses, the latter employing a novel technique to geographically cluster individuals without introducing investigator bias. The analyses revealed genetic structuring at two distinct scales. First, strikingly strong differentiation between northern and southern regions within the study area suggests little migration between them. Second, within each region, gene flow appears to be strongly limited by distance, particularly in the presence of habitat barriers such as open desert and grasslands. Northern pumas showed both reduced genetic diversity and greater divergence from a hypothetical ancestral population based on Bayesian clustering analyses, possibly reflecting a post-Pleistocene range expansion. Bayesian clustering results were sensitive to sampling density, which may complicate inference of numbers of populations when using this method. The results presented here build on those of previous studies, and begin to complete a picture of how different habitat types facilitate or impede gene flow among puma populations.     

Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels (Sciurus vulgaris) in the Italian Alps

Red squirrels (Sciurus vulgaris) are widely distributed throughout Eurasia, occurring in many types of coniferous and mixed-deciduous forests. In fragmented landscapes, small and partly isolated populations with low immigration rates show reduced genetic diversity, but reforestation can increase gene flow and restore levels of genetic variation in a few decades. No studies have so far investigated the genetic structure of red squirrel in large, continuous forests. The Italian Alps are presently characterized by almost continuous, recently reconnected forest habitats, that were affected by deep landscape changes during last glaciations but remained mostly unchanged between 10 000 and 200 years bp, when forest cover was heavily reduced. In this study we analyse patterns of genetic variability of red squirrels in and between seven sites distributed over 250 km of Alpine habitat, using mitochondrial DNA (mtDNA) and microsatellites. We use isolation-by-distance (IBD) models to investigate the relative importance that past (Pleistocene glaciations) and recent (fragmentation, bottlenecks) events had on the present genetic situation. Both nuclear and mtDNA data indicate a significant differentiation among study sites and a significant correlation between genetic and geographical distance only over a large scale. No recent bottlenecks are recorded through microsatellites and demographic models strongly support equilibrium between gene flow and drift; however, mtDNA suggests that there may have been local demographic crashes, probably in correspondence with the 19th-century forest fragmentation. These findings indicate that local landscape factors other than geographical distance per se, such as barriers of unsuitable habitat, affect gene flow and determine differentiation.     

From polar night to midnight sun: photoperiod, seal predation, and the diel vertical migrations of polar cod (Boreogadus saida) under landfast ice in the Arctic Ocean

The winter/spring vertical distributions of polar cod, copepods, and ringed seal were monitored at a 230-m station in ice-covered Franklin Bay. In daytime, polar cod of all sizes (7–95 g) formed a dense aggregation in the deep inverse thermocline (160–230 m, −1.0 to 0°C). From December (polar night) to April (18-h daylight), small polar cod <25 g migrated into the isothermal cold intermediate layer (90–150 m, −1.4°C) at night to avoid visual predation by shallow-diving immature seals. By contrast, large polar cod (25–95 g), with large livers, remained below 180 m at all times, presumably to minimize predation by deep-diving mature seals. The diel vertical migration (DVM) of small polar cod was precisely synchronized with the light/dark cycle and its duration tracked the seasonal lengthening of the photoperiod. The DVM stopped in May coincident with the midnight sun and increased schooling and feeding. We propose that foraging interference and a limited prey supply in the deep aggregation drove the upward re-distribution of small polar cod at night. The bioluminescent copepod Metridia longa could have provided the light needed by polar cod to feed on copepods in the deep aphotic layers.     

Matching genetics with oceanography: directional gene flow in a Mediterranean fish species

Genetic connectivity and geographic fragmentation are two opposing mechanisms determining the population structure of species. While the first homogenizes the genetic background across populations the second one allows their differentiation. Therefore, knowledge of processes affecting dispersal of marine organisms is crucial to understand their genetic distribution patterns and for the effective management of their populations. In this study, we use genetic analyses of eleven microsatellites in combination with oceanographic satellite and dispersal simulation data to determine distribution patterns for Serranus cabrilla, a ubiquitous demersal broadcast spawner, in the Mediterranean Sea. Pairwise population F_ST values ranged between ?0.003 and 0.135. Two genetically distinct clusters were identified, with a clear division located between the oceanographic discontinuities at the Ibiza Channel (IC) and the Almeria‐Oran Front (AOF), revealing an admixed population in between. The Balearic Front (BF) also appeared to dictate population structure. Directional gene flow on the Spanish coast was observed as S. cabrilla dispersed from west to east over the AOF, from north to south on the IC and from south of the IC towards the Balearic Islands. Correlations between genetic and oceanographic data were highly significant. Seasonal changes in current patterns and the relationship between ocean circulation patterns and spawning season may also play an important role in population structure around oceanographic fronts.     

Remnant populations of the regal fritillary (Speyeria idalia) in Pennsylvania: Local genetic structure in a high gene flow species

The Regal Fritillary butterfly, Speyeria idalia (Drury) (Lepidoptera: Nymphalidae), has been described as a high gene flow species. Supporting this assertion, previous studies in the Great Plains, where it is still relatively widespread, have found evidence of gene flow across hundreds of kilometers. Using mitochondrial and microsatellite loci, we examined the spatial genetic structure of a very isolated Pennsylvania population of these butterflies that occupies three separate meadows located within ten kilometers of each other. We found restricted gene flow and a distinct structure, with each meadow having a unique genetic signature. Our findings indicate that even a species that normally exhibits high gene flow may show fine-scale genetic subdivision in areas where populations have been largely extirpated.Authors contributed equally.     

Population subdivision and gene flow in Danish house mice

Genetic subdivision in local populations of the European house mice, Mus musculus domesticus and M. m. musculus, was analysed to study patterns of gene flow. The data consisted of frequencies of microsatellite alleles in 16 samples (250 individuals) from a total of 11 sites in Jutland, which included successive samples from three sites. Sequences of the control region of mitochondrial DNA in three successive samples from one site were also analysed. Microsatellite genotype frequencies within samples were close to Hardy-Weinberg expectations. Levels of microsatellite differentiation among samples (θ= 0.05–0.21) corresponded to limited gene flow at migration-drift equilibrium (N_m= 1–5). Weak isolation by distance for microsatellites in M. m. musculus suggested that gene flow tends to occur among neighbouring sites. Estimates of effective population size over a few generations were much lower than those corresponding to the long periods needed for arrival at mutation-drift equilibrium. This suggested that subpopulations had been influenced by gene flow since formation, or had originated recently from genetically diverse founders.     

Summer feeding ecology of harp seals (Phoca groenlandica) in relation to arctic cod (Boreogadus saida) in the Canadian high arctic

Summary The diet and feeding behaviour of harp seals, Phoca groenlandica, was examined in two high arctic locations. Fish otoliths were used to evaluate dietary composition and aspects of the population dynamics of the major prey species, arctic cod, Boreogadus saida. Harp seals, primarily adults, arrive in the high arctic in mid to late June and depart by early October. Their migration is undertaken specifically for feeding. Harp seals feed intensively on arctic cod, often occurring in dense multispecies aggregations in late summer. The average weight of harp seal stomach contents was high; glutted individuals contained as much as 6% of their body weight in food. Although arctic cod declined in abundance between years, size of cod ingested was similar between areas and years, and overlapped completely with cod taken by other marine mammals. Age/size segregation of arctic cod may account for poor representation of fish <3 years old in the seal diet. Widespread reproductive failure of arctic cod could have a profound influence on the energy balance of adult harp seals since there does not appear to be an alternate food source of equivalent energy value and abundance in arctic waters. Increasing harp seal populations will likely result in increased competition with a host of arctic cod predators, particularly ringed seals.     

Genetic diversity and gene flow within and between two different habitats of Primula merrilliana (Primulaceae), an endangered distylous forest herb in eastern China

Understanding whether and how different habitats shape population genetics is a fundamental question and a specific goal for evolutionary and conservation biology research. This study examined genetic diversity and gene flow within and between mountain and foothill habitats of Primula merrilliana, an endangered distylous forest herb in eastern China. Eleven population characteristics, including area, size and density variation, from the two habitats were also investigated. Mountain populations had significantly higher mean genetic diversity than foothill populations, which may be explained by stronger self‐incompatibility breeding system, more opportunity to use elevational shifts to track suitable sites under conditions of climate change and more heterogeneous environments in the former habitat, rather than by the differences of population size, gene flow and genetic drift intensity between them. Genetic analysis revealed that two distinct lineages, corresponding to the two habitats, diverged at China's ‘Last Glaciation’ (11 700–67 500 yr BP), suggesting this divergence was probably triggered by warmer climates during inter‐ (or post‐) glacial periods. Low unidirectional gene flow from mountain to foothill habitats, chiefly by seed dispersal, played a more important role in overall gene flow between habitats than within‐habitat gene flow. Within habitats, pollen contributes more substantially to gene flow than seed dispersal, especially in foothill habitats, possibly due to higher individual density and larger population sizes. These results have implications for the conservation in this and similar landscape areas and indicate the need to protect suitable habitats with wide elevational spans and sufficient size to permit ecological and elevational shifts in response to climatic changes. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 172–189.