Evidence of Natural Reproduction of Aquaculture-Escaped Atlantic Salmon in a Coastal British Columbia River

Abstract: We present evidence of the first successful natural spawning of Atlantic salmon (   Salmo salar ) documented on the Pacific coast of North America. Twelve juvenile Atlantic salmon composed of two year classes were captured in the Tsitika River, British Columbia. We analyzed restriction-length polymorphisms of PCR-amplified 5S rDNA and mtDNA to confirm that these individuals were Atlantic salmon. Scale analysis strongly suggested they were the products of natural spawning by feral adults. The gut contents, size, and condition of these individuals suggest that Atlantic salmon are successfully maturing in the Tsitika River, British Columbia. This event has raised concerns that the presence and possible establishment of feral Atlantic salmon may further jeopardize the continued persistence of already fragile native Pacific salmonids through competition for resources and occupation of niches that are currently underutilized.     

Assessment of harbor seal predation on adult salmonids in a Pacific Northwest estuary.

The populations of many native species have increased or expanded in distribution in recent decades, sometimes with negative consequences to sympatric native species that are rarer or less adaptable to anthropogenic changes to the environment. An example of this phenomenon from the Pacific Northwest is predation by locally abundant pinnipeds (seals and sea lions) on threatened, endangered, or otherwise depleted salmonid (Oncorhynchus spp.) populations. We used survey sampling methodology, acoustic telemetry, and molecular genetics to quantify the amount of harbor seal (Phoca vitulina) predation on a depressed run of coho salmon (O. kisutch) and to determine whether some seals consumed a disproportionately higher number of salmonids than others. Based on a probability sample totaling 759.5 h of observation, we estimated that seals consumed 1161 adult salmonids (95% CI = 503-1818 salmonids) during daylight hours over an 18.9-km estuarine study area in Oregon during an 84-d period in fall 2002. Simultaneous tracking of 56 seals via an acoustic telemetry array indicated that a small proportion of marked seals (12.5%) exhibited behavior that was consistent with specialization on salmonids. These seals spent the majority of their time in the riverine portion of the study area and did so disproportionately more at night than day. Genetic analysis of 116 salmonid structures recovered from 11 seal fecal samples suggested that coho salmon accounted for approximately one-half of total salmonid consumption. Though subject to considerable uncertainty, the combined results lead us to infer that seals consumed 21% (range = 3-63%) of the estimated prespawning population of coho salmon. We speculate that the majority of the predation occurred upriver, at night, and was done by a relatively small proportion of the local seal population. Understanding the extent and nature of pinniped predation can provide important inputs into risk assessments and other modeling efforts designed to aid the conservation and recovery of salmonids in the Pacific Northwest. Such understanding may also help inform management actions designed to reduce the impact of pinniped predation on salmonids, which potentially range from short-term lethal removal programs to long-term ecosystem restoration and protection efforts.     

Mechanistic basis of individual mortality in Pacific salmon during spawning migrations

Reproductive-based migration is a challenging period for many animals, but particularly for Pacific salmonids, which must navigate from the high seas to freshwater natal streams. For the first time, we attempt to answer the question as to why some migratory adult Pacific salmon die en route to spawning grounds. Summer-run sockeye salmon (Oncorhynchus nerka) were used as a model, and the migration behavior of 301 fish was followed by intercepting them in the ocean about 215 km from the mouth of the Fraser River, British Columbia, Canada, and implanting a gastric radio transmitter. Before release, telemetered fish were also bio-sampled, which included drawing a blood sample, collecting a gill biopsy, and quantifying energetic status with a microwave energy meter. We tested the predictions that the fish that died prematurely would be characterized by low energy reserves, advanced reproductive development, elevated indicators of stress, and low osmoregulatory preparedness compared with fish that completed their river migration. Just over half (52.3%) of the sockeye tagged were subsequently detected in the Fraser River. Salmon that failed to enter the river had exhibited indicators of stress (e.g., elevated plasma lactate, glucose, and cortisol). Contrary to our prediction, fish that failed to enter the river tended to have higher gross somatic energy and be larger at the time of sampling in the ocean than fish that successfully entered the river. Of the fish that were detected in the river (i.e., 134 fish excluding fishery removals), 9.7% did not migrate beyond the lower reaches (approximately 250 km from ocean), and a further 14.2% reached the upper reaches but failed to reach natal sub-watersheds, whereas the remainder (76.1%) reached natal sub-watersheds. Of these, fish unsuccessful in the lower reaches tended to have a high plasma osmolality in the ocean, whereas fish failing in the upper reaches had lower levels of reproductive hormones in the ocean.     

Nonrandom, Size- and Timing-Biased Breeding in a Hatchery Population of Steelhead Trout

Abstract:  Hatcheries have been built and operated to buffer salmon and trout populations from overfishing and to compensate for habitat lost or degraded by human activities. These facilities are now so prevalent that in some cases hatchery-produced salmon outnumber salmon produced in the wild. By default, this makes them an important component in the current ecology and evolution of salmonids. Hatcheries differ from natural environments in many ways, and among the most fundamental is the necessity that humans select fish for breeding instead of allowing natural processes of mate choice and competition. We examined the mating system for steelhead trout ( Oncorhynchus mykiss ) at Forks Creek Hatchery in southwest Washington and investigated factors affecting selection of individual steelhead for spawning by the hatchery staff. Despite efforts by the staff to not spawn selectively, data on steelhead spawned over 7 years revealed selection for large adult body size and early reproductive timing and a tendency for size-assortative mating (i.e., large with large). Selection on size was related to selection on reproductive timing because early returning fish tended to be larger than those returning later. To improve the fitness of both hatchery fish destined to spawn in the wild and hatchery fish designated to be spawned in the hatchery, a better understanding of factors associated with the range of reproductive success and mate-choice mechanisms in the wild is vital. This knowledge may then be applied to artificial propagation programs designed for conservation or enhancement.     

Analysis of population genetic structure of the green sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) using microsatellites

We measured within- and among-population genetic variation in the green sea urchin (Strongylocentrotus droebachiensis) at 11 sites in the north Atlantic and northeast Pacific by using four-locus microsatellite genotypes. We found no differentiation among populations from Atlantic Canada, but strong differentiation across the north Atlantic and between the Atlantic and Pacific samples. High inbreeding coefficients at three loci are consistent with high variance in reproductive success. One population that was recently decimated by disease was strongly differentiated from some others, but there was little differentiation otherwise among populations in Atlantic Canada. On a larger scale, populations in Atlantic Canada were more similar to a population from the north Pacific than to populations in the northwest Atlantic. Differentiation among populations at this large spatial scale is consistent with biogeographical hypotheses of: (1) Pleistocene population reduction and isolation in the northeast Atlantic, but (2) extinction in the northwest Atlantic followed by extensive recolonization from the Pacific. In contrast to other recent studies of trans-Atlantic organisms, we found no evidence of extensive gene flow across the north Atlantic.Communicated by R.J. Thompson, St. John's     

Population genetic structure of escolar (<Emphasis Type="Italic">Lepidocybium flavobrunneum</Emphasis>)

Escolar (Lepidocybium flavobrunneum) is a large, mesopelagic fish that inhabits tropical and temperate seas throughout the world, and is a common bycatch in pelagic longline fisheries that target tuna and swordfish. Few studies have explored the biology and natural history of escolar, and little is known regarding its population structure. To evaluate the genetic basis of population structure of escolar throughout their range, we surveyed genetic variation over an 806 base pair fragment of the mitochondrial control region. In total, 225 individuals from six geographically distant locations throughout the Atlantic (Gulf of Mexico, Brazil, South Africa) and Pacific (Ecuador, Hawaii, Australia) were analyzed. A neighbor-joining tree of haplotypes based on maximum likelihood distances revealed two highly divergent clades (δ = 4.85%) that were predominantly restricted to the Atlantic and Indo-Pacific ocean basins. All Atlantic clade individuals occurred in the Atlantic Ocean and all but four Pacific clade individuals were found in the Pacific Ocean. The four Atlantic escolar with Pacific clade haplotypes were found in the South Africa collection. The nuclear ITS-1 gene region of these four individuals was subsequently analyzed and compared to the ITS-1 gene region of four individuals from the South Africa collection with Atlantic clade haplotypes as well as four representative individuals each from the Atlantic and Pacific collections. The four South Africa escolar with Pacific mitochondrial control region haplotypes all had ITS-1 gene region sequences that clustered with the Pacific escolar, suggesting that they were recent migrants from the Indo-Pacific. Due to the high divergence and geographic separation of the Atlantic and Pacific clades, as well as reported morphological differences between Atlantic and Indo-Pacific specimens, consideration of the Atlantic and Indo-Pacific populations as separate species or subspecies may be warranted, though further study is necessary.     

Fatty acids in fish scales

This study represents the first determination of lipids and fatty acids in fish scales. Scales collected from groups of Atlantic salmon reared on fish farms and in experimental tanks were analyzed by chromatography. The complete suite of fatty acids normally found in marine organisms was detected in the scales, with the following fatty acids dominating: 16:0, 18:0, 18:1n9, 20:5n3, 22:6n3 and 24:1n9. Scales contained relatively high levels of furan fatty acids, and the level of cholesterol (2.5–5 mg/g tissue) was much higher than the levels found in the edible parts of marine fishes (0.2–1 mg/g tissue). The fatty acid profile of scales was distinct between groups of salmon originating from different commercial strains reared on the same farm, between salmon groups originating from the same strains but reared at different farms, and between groups of fed and unfed salmon in experimental tanks. Together, these data indicate that the fatty acid composition of fish scales is dependent upon both environmental and genetic factors. The fatty acid composition of fish scales may be used in stock/population identification, for example identification of escaped Atlantic salmon to farm of origin.     

Global population structure of albacore (Thunnus alalunga) inferred by RFLP analysis of the mitochondrial ATPase gene

The genetic population structure of the highly migratory albacore (Thunnus alalunga) was investigated using restriction fragment length polymorphism (RFLP) analysis of the mitochondrial ATPase gene amplified by the polymerase chain reaction (PCR). 620 individuals comprising 13 geographically distant samples (ten Pacific, two Atlantic and the Cape of Good Hope) were surveyed between 1991 and 1994 with two restriction endonucleases (Mse I and Rsa I), resulting in seven haplotypes. No heterogeneity was observed in the distribution of haplotypes among the ten samples from the North and South Pacific, nor among the samples from North and South Atlantic and Cape of Good Hope. However, highly significant heterogeneity was evident among Atlantic and Pacific samples. Higher haplotypic diversity (h) was observed in the Pacific samples (0.59 to 0.69) than in the Atlantic and Cape samples (0.22 to 0.43). These results suggest greater gene flow between albacore of the northern and southern hemispheres (within oceans) than between the Atlantic and Pacific Oceans.     

Low-level copper exposures increase visibility and vulnerability of juvenile coho salmon to cutthroat trout predators

Copper contamination in surface waters is common in watersheds with mining activities or agricultural, industrial, commercial, and residential human land uses. This widespread pollutant is neurotoxic to the chemosensory systems of fish and other aquatic species. Among Pacific salmonids (Oncorhynchus spp.), copper-induced olfactory impairment has previously been shown to disrupt behaviors reliant on a functioning sense of smell. For juvenile coho salmon (O. kisutch), this includes predator avoidance behaviors triggered by a chemical alarm cue (conspecific skin extract). However, the survival consequences of this sublethal neurobehavioral toxicity have not been explored. In the present study juvenile coho were exposed to low levels of dissolved copper (5-20 microg/L for 3 h) and then presented with cues signaling the proximity of a predator. Unexposed coho showed a sharp reduction in swimming activity in response to both conspecific skin extract and the upstream presence of a cutthroat trout predator (O. clarki clarki) previously fed juvenile coho. This alarm response was absent in prey fish that were exposed to copper. Moreover, cutthroat trout were more effective predators on copper-exposed coho during predation trials, as measured by attack latency, survival time, and capture success rate. The shift in predator-prey dynamics was similar when predators and prey were co-exposed to copper. Overall, we show that copper-exposed coho are unresponsive to their chemosensory environment, unprepared to evade nearby predators, and significantly less likely to survive an attack sequence. Our findings contribute to a growing understanding of how common environmental contaminants alter the chemical ecology of aquatic communities.     

Population structure and life history of orange roughy (Hoplostethus atlanticus) in the SW Pacific: inferences from otolith chemistry

We examined site differences in the elemental composition of the primordium and ontogenetic variability of Sr in otoliths of fish from Australia and New Zealand and, as an out-group, the North Atlantic. Differences among sites in primordium composition are slight, but significant for all five elements assayed (Sr, Pb, Cu, Zn and Hg), but principally reflect differences between the North Atlantic and SW Pacific specimens, do not replicate for independent samples in the SW Pacific and constitute a poor “natural tag” in roughy, with <25% of fish successfully assigned to source location. However, mean Sr weight-fractions at the primordium showed similar latitudinal variation across sites in Australia, New Zealand and the Tasman Sea, indicating both spatially structured populations and a common structuring process across the region. Comparisons of ontogenetic variability of Sr in otoliths from juveniles and young adults within and between sites in the SW Pacific strongly support the hypothesis that variability in this element is site-specific and environmentally sensitive, although the environmental factors involved are not obvious. The otolith analysis confirms previous suggestions that juvenile and adult Hoplostethus atlanticus are relatively sedentary, but also indicates that the population sub-structuring by age within sites is more complex and there are likely to be more spawning areas in Australian waters than previously thought. More broadly, although single point analysis of otolith composition at the primordium resolves a population structure in roughy, alone it is not precise enough to test hypotheses about the processes causing this structure. Ontogenetic variability in Sr provides better resolution of spatial structure, even in a relatively homogenous marine environment like the deep ocean, and also provides insight into behavioural and ecological factors. Ontogenetic analyses of Sr in otoliths are expensive to obtain, require more effort in specimen preparation than single point analyses, and are difficult to compare statistically, but the increased information they yield warrants their broader consideration in marine species.     

Population structure of the wreckfish Polyprion americanus determined with microsatellite genetic markers

 We examined population structure in the wreckfish, Polyprion americanus, by assaying six microsatellite loci in 422 individuals collected throughout the geographic range. Eighteen hapuku, P. oxygeneios, were assayed at the same loci for use as an outgroup. Expected heterozygosities ranged from 0.49 to 0.88 and averaged 0.66. Allele-frequency distributions at those loci indicated that samples from the eastern North Atlantic, western North Atlantic and the Mediterranean were genetically similar, confirming the pattern seen in a previous analysis of mitochondrial DNA (mtDNA) restriction fragment length polymorphisms (RFLPs). Both mtDNA and microsatellite studies differentiated northern and southern wreckfish stocks. However, in contrast to the mtDNA studies, allelic variation at microsatellite loci clearly differentiated wreckfish from two Southern Hemisphere locations, Brazil and the South Pacific. Far more genetic variation was observed at microsatellite loci than with mtDNA RFLPs (haplotype diversity averaged 0.01), and we saw more evidence of population structure with the microsatellite loci. The differentiation between southern and northern wreckfish supports the distribution records, which indicate that wreckfish do not occur in the tropics. Temperature profiles and current patterns throughout the southern oceans apparently also prevent significant gene flow between the South Pacific and Brazilian samples. Received: 29 January 2000 / Accepted: 27 June 2000     

Origin of the antitropical distribution pattern in marine mussels (Mytilus spp.): routes and timing of transequatorial migration

Many marine species, including mussels in the Mytilus edulis species group (i.e. M. edulis L., M. galloprovincialis Lamarck, and M. trossulus Gould), have an antitropical distribution pattern, with closely related taxa occurring in high latitudes of the northern and southern hemispheres but being absent from the tropics. We tested four hypotheses to explain the timing and route of transequatorial migration by species with antitropical distributions. These hypotheses yield different predictions for the phylogenetic relationship of southern hemisphere taxa relative to their northern counter-parts. The three Mytilus species were used to test these hypotheses since they exhibit a typical antitropical distribution and representative taxa occur in both the Pacific and Atlantic. Two types of mtDNA lineages were found among populations of mussels collected from the southern hemisphere between 1988 and 1996; over 90% of the mtDNA lineages formed a distinct subclade which, on average, had 1.4% divergence from haplotypes found exclusively in northern Atlantic populations of M. galloprovincialis. These data indicate that southern hemisphere mussels arose from a migration event from the northern hemisphere during the Pleistocene via an Atlantic route. The remainder of the southern hemisphere lineages (<10%) were very closely related to mtDNA haplotypes found in both M. edulis and M. galloprovincialis in the northern hemisphere, suggesting a second, more recent migration to the southern hemisphere. There was no evidence that southern hemisphere mussels arose from Pacific populations of mussels. Received: 8 December 1998 / Accepted: 8 November 1999     

Does juvenile experience affect migration and spawning of adult Atlantic salmon?

Summary This paper documents differences in seasonal time of river ascent and descent, and instream behavior of adult wild and sea-ranched Atlantic salmon (Salmo salar) of the Norwegian River Imsa stock during the period 1981–1989. Wild fish use River Imsa as a nursery, and at an age of 2 years most of them migrate to the sea as smolts. The sea ranched fish are hatchery reared offspring of the River Imsa stock and are released as smolts at the mouth of the river. They are thus deprived of juvenile river life and a downstream smolt migration. Wild and sea ranched salmon feed for 1 or more years in the Norwegian Sea before homing as spawners. Both groups returned simultaneously to coastal Norway, but sea ranched fish ascended the river later and descended sooner after spawning than wild fish. All wild females and almost all wild males (96.2%) spawned in the river, whereas 13.5% and 36.7%, respectively, of the mature sea-ranched females and males left the river unspawned. The annual number, but not the proportion, of unspawned fish increased with increasing density of adult salmon in the river. Unspawned females were medium sized and small (45–70 cm); unspawned males were medium sized and large (50–90 cm). Independent of the density of spawners in the river, sea ranched fish moved up- and downstream the river more often than wild fish. More than 20% of the sea-ranched salmon and less than 1% of the wild salmon passed a trap 100 m above the river outlet more than once in each direction during the same spawning reason. Moreover, sea-ranched salmon were about twice as often seriously injured during spawning as wild fish. Lack of juvenile experience from the river may be the main reason for the behavioral differences between sea-ranched and wild fish. Offprint requests to: B. Jonsson     

Biotic passage through the Panama Canal,with particular reference to fishes

The effectiveness of the Gatun Lake fresh-water barrier to fish migration of the Panama Canal was examined, based on literature records and recent collections. Six species have migrated to the Pacific and three to the Atlantic Oceans. New records include: Hypleurochilus aequipinnis, Barbulifer ceuthoecus and Oostethus lineatus of Atlantic origin, and Gnathanodon, speciosus of Pacific origin. The majority of Atlantic migrants are known in the Pacific only from the Miraflores Third Lock, a unique ectogenic meromictic lake attached to the Pacific entrance of the canal. The hydrography and biota (including Atlantic algal and crustacean species previously unknown as canal migrants) of the lake are discussed. It is presumed that certain fish migrants transited the canal by associating with fouling material on the underside of ships. The euryhaline species, H. aequipinnis, Lupinoblennius dispar, Lophogobius cyprinoides and Omobranchus punctatus survived in freshwater (0.0‰S) for periods longer than réquired for ship transit of the canal (ca. 8h). The stenohaline migrant Gobiosoma nudum died after 2 h in freshwater, but survived more than 50 h at 2.5‰S. Plans to increase Panama Canal ship transits through the pumping of seawater into Gatún Lake might remove the biological barrier and allow the migration of euryhaline and stenohaline species. We find those plans unwise, and the potential consequences dangerous.     

Bottom-up and climatic forcing on the worldwide population of leatherback turtles

Nesting populations of leatherback turtles (Dermochelys coriacea) in the Atlantic and western Indian Oceans are increasing or stable while those in the Pacific are declining. It has been suggested that leatherbacks in the eastern Pacific may be resource limited due to environmental variability derived from the El Ni?o Southern Oscillation (ENSO), but this has yet to be tested. Here we explored bottom-up forcing and the responding reproductive output of nesting leatherbacks worldwide. We achieved this through an extensive review of leatherback nesting and migration data and by analyzing the spatial, temporal, and quantitative nature of resources as indicated by net primary production at post-nesting female migration and foraging areas. Leatherbacks in the eastern Pacific were the smallest in body size and had the lowest reproductive output due to less productive and inconsistent resources within their migration and foraging areas. This derived from natural interannual and multidecadal climate variability together with an influence of anthropogenic climate warming that is possibly affecting these natural cycles. The reproductive output of leatherbacks in the Atlantic and western Indian Oceans was nearly twice that of turtles in the eastern Pacific. The inconsistent nature of the Pacific Ocean may also render western Pacific leatherbacks susceptible to a more variable reproductive output; however, it appears that egg harvesting on nesting beaches is their major threat. We suggest that the eastern Pacific leatherback population is more sensitive to anthropogenic mortality due to recruitment rates that are lower and more variable, thus accounting for much of the population differences compared to Atlantic and western Indian turtles.     

Biotic control of stream fluxes: spawning salmon drive nutrient and matter export

Organisms can control movements of nutrients and matter by physically modifying habitat. We examined how an ecosystem engineer, sockeye salmon (Oncorhynchus nerka), influences seasonal fluxes of sediments, nitrogen (N), and phosphorus (P) in streams of southwestern Alaska. The purpose of this study was to investigate whether salmon act as net importers or net exporters of matter and nutrients from streams and how these roles change as a function of salmon population density. We measured discharge and concentrations of suspended sediments and total N and P every 7-14 days for up to four summers in 10 streams spanning a gradient in salmon densities. We statistically allocated whole-season fluxes to salmon activities, such as excretion and bioturbation, and to export by hydrologic discharge. In addition, we used counts of spawning salmon to estimate nutrient and matter imports by salmon to streams. Large seasonal pulses of suspended sediments, P, and N were associated with salmon spawning activities, often increasing export an order of magnitude higher than during pre-salmon levels. Years and streams with more salmon had significantly higher levels of export of sediments and nutrients. In addition, years with higher precipitation had higher background export of P and N. Salmon exported an average of the equivalent of 189%, 60%, and 55% of total matter, P, and N that salmon imported in their bodies. The relative magnitude of export varied; salmon exported more than their bodies imported in 80%, 20%, and 16% across all streams and years for sediments, P, and N, respectively. A bioassay experiment indicated that the P exported by salmon is directly available for use by primary producers in the downstream lake. These results demonstrate that salmon not only move nutrients upstream on large spatial scales via their migration from the ocean and subsequent death, but also redistribute matter and nutrients on finer spatial scales through their spawning activities.     

Incorporating catastrophic risk assessments into setting conservation goals for threatened Pacific salmon

Catastrophic die-offs can have important consequences for vertebrate population growth and biodiversity, but catastrophic risks are not commonly incorporated into endangered-species recovery planning. Natural (e.g., landslides, floods) and anthropogenic (e.g., toxic leaks and spills) catastrophes pose a challenge for evolutionarily significant units (ESUs) of Pacific salmon listed under the Endangered Species Act and teetering at precariously low population levels. To spread risks among Puget Sound chinook salmon populations, recovery strategies for ESU-wide viability recommend at least two viable populations of historical life-history types in each of five geographic regions. We explored the likelihood of Puget Sound chinook salmon ESU persistence by examining spatial patterns of catastrophic risk and testing ESU viability recommendations for 22 populations of the threatened Puget Sound chinook salmon ESU. We combined geospatial information about catastrophic risks and chinook salmon distribution in Puget Sound watersheds to categorize relative catastrophic risks for each population. We then analyzed similarities in risk scores among regions and compared risk distributions among strategies: (1) population groups selected using the ESU viability recommendations of having populations spread out geographically and including historical life-history diversity, and (2) population groups selected at random. Risks from individual catastrophes varied among populations, but overall risk from catastrophes was similar within geographic regions. Recovery strategies that called for two viable populations in each of five geographic regions had lower risk than random strategies; strategies that included life-history diversity had even lower risks. Geographically distributed populations have varying catastrophic-risks profiles, thus identifying and reinforcing the spatial and life-history diversity critical for populations to respond to environmental change or needed to rescue severely depleted or extirpated populations. Recovery planning can promote viability of Pacific salmon ESUs across the landscape by incorporating catastrophic risk assessments.     

Effect of modified magnetic field on the ocean migration of maturing chum salmon, Oncorhynchus keta

To investigate the role of magnetic compass orientation in oceanic migrating chum salmon, Oncorhynchus keta, an ultrasonic telemetry study was carried out in the western North Pacific off the coast of Kushiro, Hokkaido. Four salmon were fitted with a tag which generated an artificial magnetic field and modified the geomagnetic field around the head of the fish. Initially, the free-ranging salmon with stomach-implanted ultrasonic transmitters were tracked for a period of several hours before the magnetic field was altered for a period of 16 h. The generator produced an alternating magnetic field intensity of about 6 gauss, with polarity which reversed every 11.25 min. There was no observable effect on the horizontal and vertical movements of the salmon when the magnetic field was modified. However, it was noted that salmon slowed their swimming speed significantly before changing direction, regardless of whether the fish were swimming under the normal geomagnetic field or whether they were swimming under the modified field. Received: 6 April 1997 / Accepted: 29 April 1997     

Global population divergence of the sea star Hippasteria phrygiana corresponds to the onset of the last glacial period of the Pleistocene

Genetic structure and connectivity of populations of the globally distributed and eurybathic sea star Hippasteria phrygiana (Parelius 1768) were studied in 165 individuals sampled from three oceanic regions: the North Pacific Ocean, the South Pacific Ocean (considered to include the adjacent regions of the Southern Ocean and the southern Indian Ocean) and the North Atlantic Ocean. A nuclear gene region (ATP synthase subunit α intron #5, ATPSα) and a mitochondrial gene region (cytochrome oxidase subunit I, COI) were amplified and sequenced. Significant heterogeneity was detected in an AMOVA analysis among the three sampled oceanic regions for COI, but not for ATPSα. Neither gene showed significant genetic heterogeneity within the North Atlantic, as assessed by ΦST values. Significant heterogeneity was detected for COI (but not ATPSα) in the North Pacific, but the converse was true in the South Pacific. Coalescent simulations suggested that the three regions have been diverging with little or no gene flow for the past 50–75,000 years, a time frame that corresponds to the onset of the last glacial period of the Pleistocene. A possible genetic signature of recent population expansion (or non-neutrality) was detected for each gene in the North Pacific, but not in the other two oceanic regions.     

Repeated trans-Arctic invasions in littoral bivalves: molecular zoogeography of the<Emphasis Type="Italic"> Macoma balthica</Emphasis> complex

From a geographical survey of allozyme variation, a history of repeated trans-Arctic invasions since the Plio-Pleistocene is suggested for circumboreal bivalves of the Macoma balthica complex. A principal genetic subdivision, involving several nearly diagnostic loci and Nei's distance D=0.6, distinguishes the clams of the NE Pacific from those of the NE Atlantic. The Pacific taxon is however also present in Europe, in disjunct isolates in the Baltic Sea and White Sea basins. Nevertheless, these populations have marked Atlantic introgressive elements in their gene pools (ca. 30%). Two further population types are recognized, one in the St. Lawrence estuary, Quebec, the other in Varangerfjorden, NE Norway; the latter appears a mixture of Pacific and Atlantic components in almost equal proportions, in local genetic equilibrium (a hybrid swarm). Populations in temperate North America fall outside the circumboreal M. balthica complex discussed here (D=1.0), and are referred to M. petalum. In a scenario of the history and evolution of the M. balthica complex and the similarly subdivided Mytilus edulis complex, the divergence between Pacific and Atlantic taxa started after an initial introduction of Pacific ancestors to the Atlantic basin, enabled by the Pliocene opening of the Bering Strait. During the Pleistocene and Holocene, the ocean basins were, for the most part, effectively isolated, but occasional re-invasions have taken place, causing secondary contacts of the diverged bivalve types on the Atlantic coasts. The recently re-invaded Pacific taxa in northern Europe now seem to thrive only in the extreme marginal environments. Exact dating of the re-invasions is not possible from current data. Apart from the divergence through isolation, hybridization and introgression have significantly molded the present affinities within the M. balthica complex. A formal taxonomic treatment of reticulate and hybridizing lineages is problematic; yet to recognize the evolutionary and systematic diversity within the M. balthica complex, a subspecies distinction between the NE Atlantic clams and those from the Pacific, Baltic and White Sea basins is suggested.Communicated by L. Hagerman, Helsingør