Geographical and ecological distribution of marine halacarid genera and species (Acari: Halacaridae)

At the end of 2002, the number of marine halacarid species was 1018, that of genera 51. A single genus, Copidognathus contains 33% of all species (336). Eleven genera are monotypic. Geographical provinces with a large number of species are the tropical western Pacific, temperate northeastern Atlantic, temperate southeastern Pacific, and Mediterranean-Black Sea. Most records of halacarid species are from temperate and tropical areas; 10% of species are known from polar zones. On a generic level, 29 genera are recorded from tropical and temperate but not from polar provinces, five genera are restricted to the tropics, and none to polar regions. The majority (920 species or 90%) of all species live in the upper 200 m. Records of genera with exclusively algivorous or brackish/fresh water species are bound to littoral habitats; all the other genera occur in more than one depth zone. Arenicolous genera, though most abundant in the littoral zone, have representatives in the bathyal. Four marine genera (Copidognathus, Halacarellus, Isobactrus, Lohmannella) have representatives in coastal fresh water, and three genera, Acarothrix, Caspihalacarus and Peregrinacarus, are predominantly inhabitants of diluted brackish and fresh water. None of the free-living halacarid genera of the world's oceans appears to be endemic to one geographical province.     

From Europe to America: pliocene to recent trans-atlantic expansion of cold-water north atlantic molluscs

Data on the geographical distribution, phylogeny and fossil record of cool-temperate North Atlantic shell-bearing molluscs that live in waters shallower than 100 m depth belong to two biogeographic provinces, one in eastern North America north of Cape Cod, the other in northern Europe. Amphi-Atlantic species, which are found in both provinces, comprise 30.8% of the 402 species in the northeastern Atlantic and 47.3% of the 262 species in the northwestern Atlantic. Some 54.8% of these amphi-Atlantic species have phylogenetic origins in the North Pacific. Comparisons among fossil Atlantic faunas show that amphi-Atlantic distributions became established in the Middle Pliocene (about 3.5 million years ago), and that all represent westward expansions of European taxa to North America. No American taxa spread eastward to Europe without human assistance. These results are in accord with previous phylogeographic studies among populations within several amphi-Atlantic species. Explanations for the unidirectional expansion of species across the Atlantic remain uncertain, but may include smaller size and greater prior extinction of the North American as compared to the European fauna and biased transport mechanisms. Destruction of the European source fauna may jeopardize faunas on both sides of the Atlantic.     

Global diversity of brittle stars (Echinodermata: Ophiuroidea)

This review presents a comprehensive overview of the current status regarding the global diversity of the echinoderm class Ophiuroidea, focussing on taxonomy and distribution patterns, with brief introduction to their anatomy, biology, phylogeny, and palaeontological history. A glossary of terms is provided. Species names and taxonomic decisions have been extracted from the literature and compiled in The World Ophiuroidea Database, part of the World Register of Marine Species (WoRMS). Ophiuroidea, with 2064 known species, are the largest class of Echinodermata. A table presents 16 families with numbers of genera and species. The largest are Amphiuridae (467), Ophiuridae (344 species) and Ophiacanthidae (319 species). A biogeographic analysis for all world oceans and all accepted species was performed, based on published distribution records. Approximately similar numbers of species were recorded from the shelf (n = 1313) and bathyal depth strata (1297). The Indo-Pacific region had the highest species richness overall (825 species) and at all depths. Adjacent regions were also relatively species rich, including the North Pacific (398), South Pacific (355) and Indian (316) due to the presence of many Indo-Pacific species that partially extended into these regions. A secondary region of enhanced species richness was found in the West Atlantic (335). Regions of relatively low species richness include the Arctic (73 species), East Atlantic (118), South America (124) and Antarctic (126).     

Floristic division of the Arctic

Abstract: The progress in the floristic study of the circumpolar Arctic since the 1940s is summarized and a new floristic division of this region is presented. The treeless areas of the North Atlantic and North Pacific with an oceanic climate, absence of permafrost and a very high proportion of boreal taxa are excluded from the Arctic proper. It is argued that the Arctic deserves the status of a floristic region. The tundra zone and some oceanic areas are divided into subzones according to their flora and vegetation. Two groups of subzones are recognized: the Arctic group (including the Arctic tundras proper and the High Arctic) and the Hypoarctic group. The Arctic phytochorion is floristically divided into sectors: 6 provinces and 20 subprovinces reflecting the regional features of each sector in connection with flora history, physiography and continentality-oceanity of the climate. Each sector is described and differentiated by a set of differential and co-differential species. The peculiarities of the Arctic flora are manifest in different ways in the various sectors, and endemism is not the universal criterion for subdivision.     

The biogeographical division of the Arctic and North Atlantic by the mysid (Crustacea: Mysidacea) fauna

The biogeographical analysis of Arctic and North Atlantic waters north of 30°N is based upon the distribution of 150 Mysidacea (Crustacea) species. The reasons for biogeographical divisions conducted independently by faunae of pelagic and benthopelagic mysids are adduced. The original schemes of the Arctic and North Atlantic division are proposed. Using the fauna of pelagic mysids, one biogeographical realm, one province and one transitional zone are designated in cold and temperate waters. Using the fauna of benthopelagic mysids, one biogeographical realm and four provinces are designated in those waters.     

Geographical and ecological distribution of marine halacarid genera and species (Acari: Halacaridae)

At the end of 2002, the number of marine halacarid species was 1018, that of genera 51. A single genus, Copidognathus contains 33% of all species (336). Eleven genera are monotypic. Geographical provinces with a large number of species are the tropical western Pacific, temperate northeastern Atlantic, temperate southeastern Pacific, and Mediterranean-Black Sea. Most records of halacarid species are from temperate and tropical areas; 10% of species are known from polar zones. On a generic level, 29 genera are recorded from tropical and temperate but not from polar provinces, five genera are restricted to the tropics, and none to polar regions. The majority (920 species or 90%) of all species live in the upper 200 m. Records of genera with exclusively algivorous or brackish/fresh water species are bound to littoral habitats; all the other genera occur in more than one depth zone. Arenicolous genera, though most abundant in the littoral zone, have representatives in the bathyal. Four marine genera (Copidognathus, Halacarellus, Isobactrus, Lohmannella) have representatives in coastal fresh water, and three genera, Acarothrix, Caspihalacarus and Peregrinacarus, are predominantly inhabitants of diluted brackish and fresh water. None of the free-living halacarid genera of the world's oceans appears to be endemic to one geographical province.     

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.     

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.     

Trans‐Pacific and trans‐Arctic pathways of the intertidal macroalga Fucus distichus L. reveal multiple glacial refugia and colonizations from the North Pacific to the North Atlantic

Aim We examined the phylogeography of the cold‐temperate macroalgal species Fucus distichus L., a key foundation species in rocky intertidal shores and the only Fucus species to occur naturally in both the North Pacific and the North Atlantic. Location North Pacific and North Atlantic oceans (42° to 77° N). Methods We genotyped individuals from 23 populations for a mitochondrial DNA (mtDNA) intergenic spacer (IGS) (n = 608) and the cytochrome c oxidase subunit I (COI) region (n = 276), as well as for six nuclear microsatellite loci (n = 592). Phylogeographic structure and connectivity were assessed using population genetic and phylogenetic network analyses. Results IGS mtDNA haplotype diversity was highest in the North Pacific, and divergence between Pacific haplotypes was much older than that of the single cluster of Atlantic haplotypes. Two ancestral Pacific IGS/COI clusters led to a widespread Atlantic cluster. High mtDNA and microsatellite diversities were observed in Prince William Sound, Alaska, 11 years after severe disturbance by the 1989 Exxon Valdez oil spill. Main conclusions At least two colonizations occurred from the older North Pacific populations to the North Atlantic between the opening of the Bering Strait and the onset of the Last Glacial Maximum. One colonization event was from the Japanese Archipelago/eastern Aleutians, and a second was from the Alaskan mainland around the Gulf of Alaska. Japanese populations probably arose from a single recolonization event from the eastern Aleutian Islands before the North Pacific–North Atlantic colonization. In the North Atlantic, the Last Glacial Maximum forced the species into at least two known glacial refugia: the Nova Scotia/Newfoundland (Canada) region and Andøya (northern Norway). The presence of two private haplotypes in the central Atlantic suggests the possibility of colonization from other refugia that are now too warm to support F. distichus. With the continuing decline in Arctic ice cover as a result of global climate change, renewed contact between North Pacific and North Atlantic populations of Fucus species is expected.     

TRACKING DISPERSAL ROUTES: PHYLOGEOGRAPHY OF THE ARCTIC-ANTARCTIC DISJUNCT SEAWEED ACROSIPHONIA ARCTA (CHLOROPHYTA)1

Phylogenetic relationships in the Arctic-Antarctic disjunct seaweed species Acrosiphonia arcta (Dillwyn) J. G. Agardh (Acrosiphoniales, Chlorophyta) were examined using restriction fragment-length polymorphism analysis of the fast-evolving nuclear ribosomal intergenic spacer (IGS) region and random amplified polymorphic DNA (RAPD) markers. Twenty-two isolates collected from 10 different locations in both hemispheres were compared. Five IGS length classes were identified among the 10 locations. Throughout the North Atlantic, IGS regions were found to be extremely homogeneous whereas RAPD patterns revealed subdivided populations that suggest founder effects. Acrosiphonia arcta populations found in the Arctic and North Atlantic oceans are hypothesized to be of Pacific origin. Extensive differences found between Arctic Greenland populations and those in the North Atlantic suggest that colonization of Arctic Greenland occurred as an independent event. Recolonization of the Antarctic peninsula from Southern Chile is favored, whereas the directionality of transequatorial passage along the western coast of the Americas could be in either direction.     

The discovery of new deep-sea hydrothermal vent communities in the southern ocean and implications for biogeography

Since the first discovery of deep-sea hydrothermal vents along the Galápagos Rift in 1977, numerous vent sites and endemic faunal assemblages have been found along mid-ocean ridges and back-arc basins at low to mid latitudes. These discoveries have suggested the existence of separate biogeographic provinces in the Atlantic and the North West Pacific, the existence of a province including the South West Pacific and Indian Ocean, and a separation of the North East Pacific, North East Pacific Rise, and South East Pacific Rise. The Southern Ocean is known to be a region of high deep-sea species diversity and centre of origin for the global deep-sea fauna. It has also been proposed as a gateway connecting hydrothermal vents in different oceans but is little explored because of extreme conditions. Since 2009 we have explored two segments of the East Scotia Ridge (ESR) in the Southern Ocean using a remotely operated vehicle. In each segment we located deep-sea hydrothermal vents hosting high-temperature black smokers up to 382.8°C and diffuse venting. The chemosynthetic ecosystems hosted by these vents are dominated by a new yeti crab (Kiwa n. sp.), stalked barnacles, limpets, peltospiroid gastropods, anemones, and a predatory sea star. Taxa abundant in vent ecosystems in other oceans, including polychaete worms (Siboglinidae), bathymodiolid mussels, and alvinocaridid shrimps, are absent from the ESR vents. These groups, except the Siboglinidae, possess planktotrophic larvae, rare in Antarctic marine invertebrates, suggesting that the environmental conditions of the Southern Ocean may act as a dispersal filter for vent taxa. Evidence from the distinctive fauna, the unique community structure, and multivariate analyses suggest that the Antarctic vent ecosystems represent a new vent biogeographic province. However, multivariate analyses of species present at the ESR and at other deep-sea hydrothermal vents globally indicate that vent biogeography is more complex than previously recognised.     

Australian barnacles (cirripedia: thoracica), distributions and biogeographical affinities

Currently, 279 barnacle species are recognized in Australia waters. The barnacle fauna of tropical Australia exhibits high species diversity (221), with a high incidence of tropical species (87 Indo-west Pacific [IWP], 16 West Pacific and 65 Indo-Malayan), a low species endemicity (8), and 44 cosmopolitan and 1 Australasian species. Conversely, that of temperate Australia shows lower species diversity (129), with a lower incidence of tropical species (26 IWP, 10 West Pacific and 25 Indo-Malayan), higher species endemicity (23), 37 cosmopolitan, 6 Australasian species, and 3 Australasian/Antarctic species. Distributions corroborate the general patterns demonstrated by the shallow-water biota of northern tropical and southern temperate Australian biogeographic provinces. Tropical and temperate provinces grade into each other in a broad overlap zone along both the western and eastern Australian coasts. This overlap zone is essentially a transitional region, with the gradual replacement of a tropical barnacle fauna in the north by a predominantly temperate barnacle fauna in the south. Both western and eastern Australian coasts are bounded by major poleward-flowing warm currents that have considerable influence on the marine flora and fauna, distributing tropical species of many taxa much farther south than could be predicted by latitude. Currently, 16 barnacle species introduced into Australian waters are identified, although this number may increase in the future due to new port developments and increased shipping arrivals.     

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.     

Genetic analysis of sympatric char populations in western Alaska: Arctic char (Salvelinus alpinus) and Dolly Varden (Salvelinus malma) are not two sides of the same coin

The North Pacific Ocean has been of great significance to understanding biogeography and speciation in temperate faunas, including for two species of char (Salmonidae: Salvelinus) whose evolutionary relationship has been controversial. We examined the morphology and genetics (microsatellite and mitochondrial DNA) of Arctic char (Salvelinus alpinus) and Dolly Varden char (Salvelinus malma) in lake systems in western Alaska, the eastern and western Arctic, and south of the Alaskan Peninsula. Morphologically, each lake system contained two forms: one (Arctic char) largely confined to lake habitats and characterized by greater numbers of pyloric caeca, gill rakers, and shallower bodies, and another (Dolly Varden) predominated in adjacent stream habitats and was characterized by fewer pyloric caeca, gill rakers, and deeper bodies. MtDNA partial (550 bp) d-loop sequences of both taxa were interspersed with each other within a single 'Bering' clade and demographic inferences suggested historical gene flow from Dolly Varden to Arctic char had occurred. By contrast, the taxa were strongly differentiated in sympatry across nine microsatellite loci in both lakes. Our data show that the two taxa are highly genetically distinct in sympatry, supporting their status as valid biological species, despite occasional hybridization. The interaction between these species highlights the importance of the North Pacific, and Beringia in particular, as an evolutionary wellspring of biodiversity.     

Molecular lineage diversity and inter‐oceanic biogeographical history in Hiatella (Mollusca,Bivalvia)

Hiatella is one of the most widespread marine bivalve genera, occurring in diverse habitats from the temperate to polar latitudes in both hemispheres, and in fossil strata since almost 150 Myr ago. Despite variation in some biological and morphological traits, characters to resolve the current systematic structure consistently across the range of the genus are not known: all samples are often referred to a single species, Hiatella arctica (L.). Exploring the systematics of Hiatella using partial sequences of three genes (mitochondrial COI, and the nuclear ANT and 28S rRNA), we find high diversity of deep lineages (11–22% p‐distance in COI), and identify at least 13 distinct taxa both by heuristic criteria (congruence of the nuclear and mtDNA data) and by coalescence‐based analyses. At several localities, two or three of these cryptic species were found in sympatry. In the framework of previous fossil evidence and of hypotheses of paleoceanographical connections, scenarios of the phylogeny and biogeographical history of the identified species at a range of different time scales are outlined. The distinction between the main North Pacific and North Atlantic Hiatella clades and systematic diversification within each of them seem to have followed a Miocene trans‐Panamanian invasion. Apart from such earlier intra‐basin diversification, the data suggest that three successive counter‐invasions from the Pacific to the Atlantic via the Arctic Ocean route have later contributed to the current North Atlantic Hiatella diversity. These invasions probably took place in connection with (i) the Great Trans‐Arctic Biotic Interchange in the Pliocene, (ii) the last interglacial period c. 120 kya and (iii) the Holocene, postdating the last glaciation. This sequence of trans‐Arctic invasions is largely analogous to that hypothesized for some other boreal‐arctic bivalves.     

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.     

Low colonisation on artificial substrata in arctic Spitsbergen

High polar communities tend to be young because of the frequent and intense impact of ice (scour), so colonisation patterns are particularly important. Yet, despite a wealth of studies at temperate and tropical latitudes, we know of no hard substratum settlement/colonisation experiments reported north of 60°N, to date. Here we report on fauna encrusting square panels immersed at 12 m depth in Isforden, Spitsbergen (Svalbard) after 2 and 3 days, a week and a year. Arctic colonisation is slow but is not species poor. We found no colonists present after 2 and 3 days but two panels were colonised by a bryozoan and polychaetes after a week. After a year immersion, three panels were 3, 5 and 11% covered with a mean of ∼247 colonists. This is about an order of magnitude lower than has been described from most studies elsewhere, but not as low as has been recorded at an Antarctic locality. Most individual colonists (80–93%) were polychaetes (Spirorbis tridentatus) but most of the species were bryozoans. The Arctic is widely described as taxon poor compared with elsewhere, but at the local scale we investigated, species richness (20) was as high or higher than in many similar colonisation studies along the north Pacific or Atlantic coasts. In striking contrast, no settlement panel study has yielded fewer higher taxa (2 phyla, 3 classes) than this high arctic study.     

Multiple fossil calibrations,nuclear loci and mitochondrial genomes provide new insight into biogeography and divergence timing for true seals (Phocidae,Pinnipedia)

Aim To better understand the historical biogeography of the true seals, Phocidae, by combining nuclear DNA (nDNA) and mitochondrial DNA (mtDNA) in a divergence time analysis using multiple fossil calibrations. Location Arctic, Antarctic, Pacific and Atlantic Oceans, Lake Baikal, Caspian Sea. Methods Fifteen nuclear genes totalling 8935 bp plus near‐complete mitochondrial genome sequences were used in a Bayesian divergence time analysis, incorporating eight soft‐bound fossil calibrations across the phylogeny. All species of true seals were included, plus the walrus, three otariids and seven carnivore outgroups. The majority of the nuclear sequences and four phocid mitochondrial genomes (plus three non‐phocid mitochondrial genomes) were newly generated for this study using DNA extracted from tissue samples; other sequences were obtained from GenBank. Results Using multiple nuclear genes and multiple fossil calibrations resulted in most divergence time estimations within Phocidae being much more recent than predicted by other molecular studies incorporating only mtDNA and using a single calibration point. A new phylogenetic hypothesis was recovered for the Antarctic seals. Main conclusions Incorporating multiple nuclear genes and fossil calibrations had a profound effect on the estimated divergence times. Most estimated divergences within Phocinae (Arctic seals) correspond to Arctic oceanic events and all occur within the last 12 Myr, a time when the Arctic and Atlantic oceans were freely exchanging and perennial Arctic sea ice existed, indicating that the Arctic seals may have had a longer association with ice than previously thought. The Monachinae (‘southern’ seals) split from the Phocinae c. 15 Ma on the eastern US coast. Several early trans‐Atlantic dispersals possibly occurred, leaving no living descendants, as divergence estimates suggest that the Monachus (monk seal) species divergences occurred in the western Atlantic c. 6 Ma, with the Mediterranean monk seal ancestor dispersing afterwards. The tribes Lobodontini (Antarctic seals) and Miroungini (elephant seals) are also estimated to have diverged in the eastern Atlantic c. 7 Ma and a single Lobodontini dispersal to Antarctica occurred shortly afterwards. Many of the newly estimated dates are used to infer how extinct lineages/taxa are allied with their living relatives.