The Bering Strait connection: dispersal and speciation in boreal macroalgae

A large number of boreal seaweeds have either sibling species or conspecific populations of a single species in the North Pacific and North Atlantic Oceans. This pattern is thought to have arisen from the dispersal between the two oceans through the Arctic Ocean after the opening of the Bering Strait in the mid-to-late Miocene or earliest Pliocene and from subsequent vicariant speciation as the Arctic Ocean froze and Bering Strait closed intermittently during glacial periods. Recent molecular studies of species in all three major seaweed phyla reveal patterns of vicariance. However, a number of lines of evidence point to differences in origins of these clades; some appear to be Pacific in origin whereas others appear to be derived from Atlantic stock. Different origins can be explained by recent stratigraphic finds that push the first Cenozoic opening of the Bering Strait back from 3.1–4.1 to 4.8–7.4 Ma (million years ago). Northern hemisphere ocean circulation models suggest that water flow would have been from the North Atlantic–Arctic south through the Bering Strait prior to the closure of the Panamanian Isthmus c. 3.5 Ma in contrast to the northward flow from the Pacific into the Arctic and North Atlantic, which developed after the closing of the Isthmus. Despite these differences in timing of the two invasions, there are no significant differences in levels of relationships among species with a North Atlantic origin compared with species with a North Pacific origin based on currently available data. More work is required to understand vicariance in seaweeds, especially in deciphering when a speciation event has occurred.     

THE BIOGEOGRAPHIC ORIGIN OF ARCTIC ENDEMIC SEAWEEDS: A THERMOGEOGRAPHIC VIEW1

The Arctic is geologically and biogeographically young, and the origin of its seaweed flora has been widely debated. The Arctic littoral biogeographic region dates from the latest Tertiary and Pleistocene. Following the opening of Bering Strait, about 3.5 mya, the “Great Trans‐Arctic Biotic Interchange” populated the Arctic with a fauna strongly dominated by species of North Pacific origin. The Thermogeographic Model (TM) demonstrates why climate and geography continued to support this pattern in the Pleistocene. Thus, Arctic and Atlantic subarctic species of seaweeds are likely to be evolutionarily “based” in the North Pacific, subarctic species are likely to be widespread in the warmer Arctic, and species of Atlantic Boreal or warmer origin are unlikely in the Arctic and Subarctic. Although Arctic seaweeds have been thought to have a greater affinity with the North Atlantic, we have reanalyzed the Arctic endemic algal flora, using the Thermogeographic Model and evolutionary trees based on molecular data, to demonstrate otherwise. There are 35 congeneric species of the six, abundant Arctic Rhodophyta that we treat in this paper; 32 of these species (91%) occur in the North Pacific, two species (6%) occur in the Boreal or warmer Atlantic Ocean, and a single species is panoceanic, but restricted to the Subarctic. Laminaria solidungula J. Agardh, a kelp Arctic “endemic” species, has 18 sister species. While only eleven (61%) occur in the North Pacific, this rapidly dispersing and evolving genus is a terminal member of a diverse family and order (Laminariales) widely accepted to have evolved in the North Pacific. Thus, both the physical/time‐based TM and the dominant biogeographic pattern of relatives of Arctic macrophytes suggest strong compliance with the evidence of zoology, geology, and paleoclimatology that the Arctic marine flora is largely of Pacific origin.     

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.     

2010年夏季白令海小型浮游植物分布

根据2010年7月10-19日我国第四次北极科学考察“雪龙”号考察船在白令海(52°42.29′-65°30.23′ N, 169°20.85′ E-179°30.37′ W)采集的70份水采样品,共鉴定小型浮游植物5个门类143种(含变种和变型).其中硅藻门37属95种,甲藻门15属44种,绿藻门2属2种,裸藻门和金藻门各1属1种.聚类分析表明: 调查海区浮游植物可分为深水区群落和浅水区群落.深水区群落分布于太平洋西北部和白令海海盆,种类组成主要以温带大洋性种西氏新细齿状藻、大西洋角毛藻和广布种菱形海线藻、扁面角毛藻为主,浮游植物的丰度较低,种间分配均匀,优势种不突出,种类多样性指数高;浅水区群落分布于白令海陆坡区和北部陆架区,主要由近岸冷水种诺登海链藻、叉尖角毛藻和广温广盐种丹麦细柱藻、旋链角毛藻等组成,浮游植物的丰度高,种间分配不均匀,优势种突出,种类多样性指数低.浮游植物平均丰度为58722 cells·L^-1,变化范围在950～192400 cells·L^-1,站间差异显著.平面分布趋势总体呈白令海陆架区＞白令海陆坡区＞白令海海盆＞太平洋西北部海域.垂直分布均以表层浮游植物丰度较低,至温跃层附近出现高值.不同水域温跃层的差异决定了其垂直分布格局.     

Composition of the summer photosynthetic pico and nanoplankton communities in the Beaufort Sea assessed by T-RFLP and sequences of the 18S rRNA gene from flow cytometry sorted samples

The composition of photosynthetic pico and nanoeukaryotes was investigated in the North East Pacific and the Arctic Ocean with special emphasis on the Beaufort Sea during the MALINA cruise in summer 2009. Photosynthetic populations were sorted using flow cytometry based on their size and pigment fluorescence. Diversity of the sorted photosynthetic eukaryotes was determined using terminal-restriction fragment length polymorphism analysis and cloning/sequencing of the 18S ribosomal RNA gene. Picoplankton was dominated by Mamiellophyceae, a class of small green algae previously included in the prasinophytes: in the North East Pacific, the contribution of an Arctic Micromonas ecotype increased steadily northward becoming the only taxon occurring at most stations throughout the Beaufort Sea. In contrast, nanoplankton was more diverse: North Pacific stations were dominated by Pseudo-nitzschia sp. whereas those in the Beaufort Sea were dominated by two distinct Chaetoceros species as well as by Chrysophyceae, Pelagophyceae and Chrysochromulina spp.. This study confirms the importance of Arctic Micromonas within picoplankton throughout the Beaufort Sea and demonstrates that the photosynthetic picoeukaryote community in the Arctic is much less diverse than at lower latitudes. Moreover, in contrast to what occurs in warmer waters, most of the key pico- and nanoplankton species found in the Beaufort Sea could be successfully established in culture.     

Phylogeny of the eelpout genus Lycodes (Pisces,Zoarcidae) as inferred from mitochondrial cytochrome b and 12S rDNA

The bottom-dwelling and species-rich eelpout genus Lycodes Reinhardt has a great potential for the study of Arctic marine speciation. Subdivision of the genus has been based on single or few morphological characters (e.g., lateral line configuration) with contradicting results and phylogenetic approaches have not been attended. Here we present the first phylogenetic analysis of the genus employing DNA sequences of the mitochondrial genes cytochrome b and 12S rDNA (714 bp). The analysis with the two genes combined resulted in two equally parsimonious trees. In both cladograms most of the previously suggested subgroups are para- or polyphyletic, except for the so-called short-tailed Lycodes spp., with a short tail, a single mediolateral lateral line configuration and a shallow or filled otolith sulcus. The group of long-tailed Lycodes spp., with ventral or ventro-medio-lateral types of lateral line configuration and a deep otolith sulcus, appears to be paraphyletic, since Pacific and Atlantic species in this group are not each other's closest relatives. Thus, the short-tailed species are placed in a derived clade, indicating a secondary shortening of the tail, and a "slope to shore" type of evolution. This is not in accordance with earlier assumptions of the more elongate, deeper living species being the more derived. The basal position of long-tailed Pacific species supports earlier theories of Pacific origin of the genus/family. Small genetic differences between Arctic/Atlantic species indicate a rather recent radiation in these areas after the opening of the Bering Strait 3.0-3.5 million years ago.     

Biogeography of the benthic marine algae of the North Atlantic Ocean—an overview

An overview of the biogeography of the benthic marine algae of the North Atlantic Ocean is presented. General and specific distribution patterns are discussed in the light of current knowledge of extant species, and of known events in the evolution of the North Atlantic Ocean. The close relationships between the Arctic, NW and NE Atlantic floras suggest their possible origin as a single flora in the early Oligocene Arctic Ocean, when it was isolated by the Bering Land Bridge and the Greenland-Scotland Ridge. Migration of the flora into the North Atlantic Ocean could have occurred with the subsidence of the Greenland-Scotland Ridge. The present day distribution patterns are the main clue to unravelling the past, and study of vicariant amphi-Atlantic taxa using a variety of experimental techniques will yield the most valuable information in attempts to interpret major biogeographical events in the North Atlantic Ocean.     

Population status of Greenland halibut <Emphasis Type="Italic">Reinhardtius hippoglossoides</Emphasis> (Walbaum, 1793) of the Laptev Sea

This is the first study to perform a comparative genetic analysis of Greenland halibut in the samples from the Atlantic (waters of west and east of Greenland), Arctic (Laptev Sea), and Pacific (the western part of the Bering Sea) ocean basins using seven microsatellite loci. The obtained data clearly demonstrate that the Greenland halibut population in the Laptev Sea belongs to the groups of the Atlantic Ocean basin. Apparently, the Greenland halibut of the Laptev Sea is represented by a dependent population, which is replenished due to the drift of immatures from the spawning grounds in the Barents Sea with the transformed Atlantic water flow along the continental slope. In addition, the Arctic population can be partially replenished due to the breeding of the halibut in local spawning grounds.     

DNA evidence for a Paleocene origin of the Alcidae (Aves: Charadriiformes) in the Pacific and multiple dispersals across northern oceans

The Alcidae is a group of marine, wing-propelled diving birds known as auks that are distributed along the coasts of the northern oceans. It has been suggested that auks originated in the Pacific coastal shores as early as the Miocene, and dispersed to the Atlantic either through the Arctic coasts of Eurasia and North America (northern dispersal route), or through upwelling zones in the coastal areas of California to Florida (southern dispersal route), before the closure of the Isthmus of Panama in the Pliocene. These hypotheses have not been tested formally because proposed phylogenies failed to recover fully bifurcating, well-supported phylogenetic relationships among and within genera. We therefore constructed a large data set of mitochondrial and nuclear DNA sequences for 21 of the 23 species of extant auks. We also included sequences from two other extant and one extinct species retrieved from GenBank. Our analyses recovered a well-supported phylogenetic hypothesis among and within genera. Aethia is the only genus for which we could not obtain strong support for species relationships, probably due to incomplete lineage sorting. By applying a Bayesian method of molecular dating that allows for rate variation across lineages and genes, we showed that auks became an independent lineage in the Early Paleocene and radiated gradually from the Early Eocene to the Quaternary. Reconstruction of ancestral areas strongly suggests that auks originated in the Pacific during the Paleocene. The southern dispersal route seems to have favored the subsequent colonization of the northern Atlantic Ocean during the Eocene and Oligocene. The northern route across the Arctic Ocean was probably only used more recently after the opening of the Norwegian Sea in the Middle Miocene and the opening of the Bering Strait in the Late Miocene. We postulate that the ancestors of auks lived in a warmer world than that currently occupied by auks, and became gradually adapted to feeding in cool marine currents with high biomass productivity. Hence, warmer tropical waters are now a barrier for the dispersal of auks into the Southern Hemisphere, as it is for penguins in the opposite direction.     

Genetic Stock Identification of Chum Salmon in the Bering Sea and North Pacific Ocean Using Mitochondrial DNA Microarray

A newly developed DNA microarray was applied to identify mitochondrial (mt) DNA haplotypes of more than 2200 chum salmon in the Bering Sea and North Pacific Ocean in September 2002 and also 2003, when the majority of maturing fish were migrating toward their natal river. The distribution of haplotypes occurring in Asian and North American fish in the surveyed area was similar in the 2 years. A conditional maximum likelihood method for estimation of stock compositions indicated that the Japanese stocks were distributed mainly in the north central Bering Sea, whereas the Russian stocks were mainly in the western Bering Sea. The North American stocks were abundant in the North Pacific Ocean around the Aleutian Islands. These results indicate that the Asian and North American stocks of chum salmon are nonrandomly distributed in the Bering Sea and the North Pacific Ocean, and further the oligonuleotide DNA microarray developed by us has a high potential for identification of stocks among mixed ocean aggregates of high-seas chum salmon.     

A Review of the Sipunculan Genus Thysanocardia Fisher

The taxonomy of the sipunculan taxon Thysanocardia Fisher is reviewed. Originally erected as a subgenus of Golfingia Lankester, the status of Thysanocardia is here elevated to the level of a genus. Of the 17 species currently recognised within this genus, most of which were originally described from single specimens, only three are considered sufficiently distinct as to warrant specific status, namely T. catherinae (Grube), T. procera (Möbius) and T. nigra (Ikeda). T. catherinae is the most widespread of these species being distributed in the western North Atlantic, South Atlantic and western Indian Ocean, whilst T. procera is confined to the north-eastem region of the North Atlantic and T. nigra chiefly to the North Pacific Ocean.     

Evolution of the Northern Rockweed,Fucus distichus,in a Regime of Glacial Cycling: Implications for Benthic Algal Phylogenetics

Northern hemisphere rockweeds (Fucus) are thought to have evolved in the North Pacific and then spread to the North Atlantic following the opening of the Bering Strait. They have dispersed and widely speciated in the North Atlantic and its tributary seas. Fucus distichus is likely near the ancestral member of this genus, and studies have shown that there are several species/subspecies in this complex (i.e. F. evanescens and F. gardneri). We used phylogenetic and haplotype analyses to test the phylogenetic relationships and biogeography of F. distichus. Our data and subsequent analyses demonstrate that, unlike previous studies that lacked samples from an extensive geographical area of the Arctic and Subarctic, there is a distinct Arctic haplotype that is the source of subspecies in both the North Pacific and North Atlantic. Fucus distichus occupies a low tide zone habitat, and in Arctic/Subarctic regions it is adapted to the severe stress of sea ice coverage and disturbance during many months per year. We hypothesize that the very large geographic area of Arctic and Subarctic rocky shores available to this species during interglacials, supported by large Arctic/Subarctic fringe areas as well as unglaciated refugia during glacial cycles, provided a robust population and gene pool (described by the Thermogeographic Model). This gene pool dilutes that of the more fragmented and area-limited Temperate/Boreal area populations when they are brought together during glacial cycles. We suggest that similar subspecies complexes for a variety of Arctic/Subarctic shore biota should be examined further in this context, rather than arbitrarily being split up into numerous species.     

Systematic review of the genus <Emphasis Type="Italic">Bothrocara</Emphasis> Bean 1890 (Teleostei: Zoarcidae)

The systematics of the eelpout genus Bothrocara Bean 1890 is reviewed on the basis of 941 specimens. Eight mostly eurybathic, demersal species are recognized, distributed mainly along the continental slopes of the North and South Pacific oceans, with one species entering the South Atlantic. Distributions are: B. brunneum ranges from the Sea of Okhotsk to the Gulf of Panama at depths of 199–1,829 m; B. elongatum ranges from the Gulf of Panama to Chile at depths of 720–1,866 m; B. hollandi ranges from the Sea of Japan to the southeastern Bering Sea at depths of 150–1,980 m; B. molle ranges from the western Bering Sea to the South Atlantic at depths of 106–2,688 m; B. nyx is known only from the eastern Bering Sea at depths of 790–1,508 m; B. pusillum ranges from the northern Bering Sea to British Columbia, Canada, at depths of 55–642 m; B. tanakae is found along the northern coasts of Honshu and Hokkaido islands, Japan, at depths of 274–892 m; B. zestum ranges from the Izu Islands, Japan, and central Honshu, Japan, to the Gulf of Alaska at depths of 199–1,620 m (an unidentifiable specimen from off Taiwan may be B. zestum). The species are distinguished from one another mainly on the basis of head pore patterns, gill raker morphology, coloration and various meristic and morphometric values. A determination key to the species is provided.     

A mitogenomic approach to the taxonomy of pollocks: <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> and <Emphasis Type="Italic">T. finnmarchica</Emphasis> represent one single species

Background  

The walleye pollock (Theragra chalcogramma) and Norwegian pollock (T. finnmarchica) are confined to the North Pacific and North Atlantic Oceans, respectively, and considered as distinct species within the family Gadidae. We have determined the complete mtDNA nucleotide sequence of two specimens of Norwegian pollock and compared the sequences to that of 10 specimens of walleye pollock representing stocks from the Sea of Japan and the Bering Sea, 2 specimens of Atlantic cod (Gadus morhua), and 2 specimens of haddock (Melanogrammus aeglefinus).     

Evidence for initial opening of the Pacific ocean in the Jurassic

An argument is presented for the east-west initial opening of the pacific Ocean during Early Jurassic times, on paleontological, biogeograhical, and geological grounds. A new reconstruction of the continents bordering the Pacific Ocean is proposed, based chiefly upon trans-Pacific, terrestrial biotic links that are not reflected in regions bordering the Atlantic. A new Arctic Ocean reconstruction is also proposed. Since the Atlantic and Indian Oceans were closed in the Early Jurassic as well, an expanding-earth model is adopted, instead of a Pangaen reconstruction on a present-sized earth. The model prposed here is also compatible with geological evidence for a narrow, epicontinental Tethys Sea. Sea-floor subduction along the perimeter of the Pacific appears to be actively taking place and in the past was much more extensive along the eastern border than on the western side. The tectonics of Antarctica, the Southwest Pacific, Indonesia, Japan, the Arctic region, western North America, Baja California, the Carribean, and the Gulf of Mexico are discussed in terms of the proposed model. The Pacific sea-floor spreading history is outlined and related to the initial opening of the North Atlantic 180 million years ago (Early Jurassic). The South Pacific and Indian Ocean are both considered to have opened some 155 million years ago (Late Jurassic). The original and present positions of the Jurassic equator are determined.     

Origin, diversification, and historical biogeography of the genus Trachurus (Perciformes: Carangidae)

We addressed phylogenetic relationships in the genus Trachurus using cytochrome b gene and D-loop sequences. The trees showed five groups: (1) the Southwest Pacific species (T. japonicus, T. novaezelandiae, and T. declivis); (2) The Mediterranean Sea and Eastern Atlantic species (T. mediterraneus); (3) The Atlantic Ocean species (T. lathami and T. trecae); (4) Eastern Atlantic species (T. trachurus and T. capensis); and (5) a group of highly mobile pelagic species, two from the Eastern Pacific (T. symmetricus and T. murphyi) and one from the Eastern Atlantic (T. picturatus). The phylogeny based on Cyt b, supports the molecular clock hypothesis and our results agree with the reported fossil indicating that the origin of this genus occur when the Thetys Sea closed (around 18.4 MYA). In addition, a very slow neutral substitution rate is reported identified only two periods of maximum diversification: the first occurring between 18.4 and 15.0 MYA and the second between 8.4 MYA and present day.     

A biological consequence of reducing Arctic ice cover: arrival of the Pacific diatom Neodenticula seminae in the North Atlantic for the first time in 800 000 years

The Continuous Plankton Recorder survey has monitored plankton in the Northwest Atlantic at monthly intervals since 1962, with an interegnum between 1978 and 1990. In May 1999, large numbers of the Pacific diatom Neodenticula seminae were found in Continuous Plankton Recorder (CPR) samples in the Labrador Sea as the first record in the North Atlantic for more than 800 000 years. The event coincided with modifications in Arctic hydrography and circulation, increased flows of Pacific water into the Northwest Atlantic and in the previous year the exceptional occurrence of extensive ice‐free water to the North of Canada. These observations indicate that N. seminae was carried in a pulse of Pacific water in 1998/early 1999 via the Canadian Arctic Archipelago and/or Fram Strait. The species occurred previously in the North Atlantic during the Pleistocene from∼1.2 to∼0.8 Ma as recorded in deep sea sediment cores. The reappearance of N. seminae in the North Atlantic is an indicator of the scale and speed of changes that are taking place in the Arctic and North Atlantic oceans as a consequence of regional climate warming. Because of the unusual nature of the event it appears that a threshold has been passed, marking a change in the circulation between the North Pacific and North Atlantic Oceans via the Arctic. Trans‐Arctic migrations from the Pacific into the Atlantic are likely to occur increasingly over the next 100 years as Arctic ice continues to melt affecting Atlantic biodiversity and the biological pump with consequent feedbacks to the carbon cycle.