Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Evidence of exposure to Brucella sp. in harbor porpoises (Phocoena phocoena) from the Bay of Fundy, Canada

Novel strains of Brucella recently have been discovered in marine mammals. To investigate Brucella exposure and infection in a general population of cetaceans, blood and tissue samples were collected and analyzed from wild harbor porpoises (Phocoena phocoena) incidentally caught in fishing gear in the Bay of Fundy, Canada. Two of 170 (1.2%) animals had detectable antibodies against Brucella, but no organisms were isolated from genital swabs or tissues from 22 and 8 porpoises, respectively. Genetic analysis of inflamed testes from 20 animals yielded no amplification of Brucella DNA. This is the first evidence of exposure to Brucella in porpoises from the western North Atlantic, and the prevalence is much lower than documented for conspecifics from the eastern North Atlantic.     

Trends in the maternal investment of harbour porpoises are uncoupled from the dynamics of their primary prey

Harbour porpoises in the Bay of Fundy and Gulf of Maine feed primarily on Atlantic herring. Herring stocks have undergone dramatic fluctuations in abundance over the past three decades due to changes in fishing intensity. In order to understand the effects of these changes in prey abundance on the patterns of maternal investment, I examined interdecadal variation in the size of porpoise calves measured in August prior to weaning. Female harbour porpoises exhibited significant variation in maternal investment between 1970 and 1999. During the 1980s, females consumed more herring and produced larger calves. Surprisingly, however, this increased maternal investment occurred during the period of lowest prey abundance, perhaps because the herring stock assessment does not reflect the availability or quality of prey to female porpoises.     

BLOOD CHEMISTRY OF WILD HARBOR PORPOISES PHOCOENA PHOCOENA (L.)

Blood chemistry values were measured from 31 harbor porpoises ( Phocoena phocoena ) released from herring weirs in the Bay of Fundy, Canada. Sodium, phosphorus, calcium, chloride, magnesium, total protein, albumin, globulin, urea, cholesterol, serum osmolality, and alanine aminotransferase levels fell within ranges reported for captive harbor porpoises and other odontocetes. Glucose, potassium, creatine kinase, aspartate aminotransferase, hemoglobin, thyroxine, bilirubin, and alkaline phosphatase levels were generally higher than those reported for captive odontocetes. Cortisol and creatinine levels were significantly higher ( P = 0.0356 and 0.0174, respectively) in porpoises that were handled for longer periods and fitted with electronic tags (mean cortisol value = 314 ± 107 nmol/L, mean creatinine value = 94 ± 14 μmol/L) than those receiving roto-tags (mean cortisol value = 222 ± 82 nmol/L, mean creatinine value = 75 ± 15 μmol/L). A more controlled study of duration of handling, time spent out of water, degree of manipulation, and method of tag attachment is required to identify which factors are responsible for the stress response.     

Microsatellite loci reveal highly significant genetic differentiation among Atlantic salmon (Salmo salar L.) stocks from the east coast of Canada

Allele frequency data from eight microsatellite loci provide evidence of highly significant genetic differentiation among stocks of Atlantic salmon Salmo salar L. from the Bay of Fundy, eastern and north-western Nova Scotia and Newfoundland. Estimates of genetic structure ( R _ST and θ) were significant both among all samples taken from the different geographical locations and among samples from geographical regions for which more than one stock was sampled. Samples from the Bay of Fundy taken from stocks which are phenotypically and behaviourally diverse showed particularly high levels of genetic structure. Rogers', allele sharing and (δμ)² distances also revealed significant differences among stock samples and were significantly correlated [Rogers' and (δμ)²] with sea distance between rivers. Results suggest that stocks of Atlantic salmon in eastern Canada are highly diverse genetically and that this should be an important consideration in any management programme for stocks in the area.     

HEMATOLOGY VALUES OF WILD HARBOR PORPOISES (PHOCOENA PHOCOENA) FROM THE BAY OF FUNDY,CANADA

Clinical hematology values were determined for 29 harbor porpoises (Phocoena phocoena) released from herring weirs in the Bay of Fundy, Canada. Erythrocyte values exhibited narrow ranges, but there was a high degree of individual variability in counts of white blood cells. Total white cell counts ranged from 2.6 to 15.5 ± 10⁹/liter, with an overall mean of 6.5 ± 2.7 ± 10±/liter. There were significant differences among reproductive classes in mean values of total red blood cells, hematocrit (HCT), mean cell volume (MCV), mean cell hemoglobin (MCHC) and total monocyte count. Wild harbor porpoises had fewer white blood cells, lower relative total number of neutrophils and lymphocytes, and higher percentages of monocytes and eosinophils than reported in the literature for captive porpoises. Compared to published values for other odontocetes, the hemograms of harbor porpoises were most similar to those of Pacific white-sided dolphins (Lagenorhynchus acutus). These hematology data represent a baseline from free-ranging harbor porpoises that can be used as a reference for long-term monitoring of the health of this population and as a tool for rehabilitation facilities.     

LIFE IN THE FAST LANE: THE LIFE HISTORY OF HARBOR PORPOISES FROM THE GULF OF MAINE

We describe the life history of harbor porpoises in the Gulf of Maine by examining 239 animals killed in gill net fisheries and comparing these findings with the results of previous studies from the Bay of Fundy. Most female porpoises matured at age three and became pregnant each year thereafter. Reproduction was strictly seasonal, with ovulation, conception, and parturition occurring in the spring and early summer. The oldest specimen in the sample was 17 yr of age, but most individuals were younger than 12. The findings are similar to those of earlier studies from the Bay of Fundy and support the hypothesis that these animals form a single population. Harbor porpoises represent one end of a continuum of odontocete life histories that spans a wide diversity of strategies. In comparison with other, large odontocetes, harbor porpoises mature at an earlier age, reproduce more frequently, and live for shorter periods.     

Life-stage-dependent supergene haplotype frequencies and metapopulation neutral genetic patterns of Atlantic cod,Gadus morhua,from Canada's Northern cod stock region and adjacent areas

Among highly migratory fish species, nursery areas occupied by juveniles often differ from adult habitats. To better understand the spatial dynamics of Canada's Northern cod stock, juveniles caught off the east coast of Newfoundland and Labrador were compared to adults from the same region as well as individuals from other areas in Atlantic Canada using double-digest restriction site-associated DNA sequencing–derived single nucleotide polymorphisms. A reduced proportion of homozygotes with a chromosomal inversion located in linkage group 1 (LG1) was detected between juvenile and adult samples in the Northern cod stock region, potentially indicating age-dependent habitat use or ontogenetic selection for attributes associated with the many genes located in LG1. No selectively neutral genetic differences were found between samples from the Northern cod stock; nevertheless, significant differences were found between some of these samples and cod collected from St. Pierre Bank, Bay of Fundy, Browns Bank and the southern Scotian Shelf. Clustering analysis of variants at neutral loci provided evidence for three major genetic units: (a) the Newfoundland Atlantic Coast, (b) eastern and southern Gulf of St. Lawrence and Burgeo Bank and (c) the Bay of Fundy, Browns Bank and southern Scotian Shelf. Both adaptive and neutral population structure within the Northern cod stock should be considered by managers to promote demographic rebuilding of the stock.     

Macrotidal estuaries: a region of collision between migratory marine animals and tidal power development

Macrotidal estuaries of the inner Bay of Fundy are utilized by large numbers of migratory fishes, particularly dogfish, sturgeon, herring, shad, Atlantic salmon and striped bass as well as by other migratory marine animals, many of which have large body sizes (squid, Lamnid sharks, seals and whales). Tagging experiments indicate the fishes originate from stocks derived over the entire North American Atlantic coast from Florida to Labrador. Population estimates suggest up to 2.0 times 10⁶adult American shad (Alosa sapidissima) migrate through an individual embayment each year. These migrations are an integral part of the life history of the respective species and appear to be controlled in part by the near shore movements of ocean currents. In other regions of the world similar macrotidal estuaries exist (Cook Inlet, Alaska; Severn Estuary, U.K.) and they, like the Bay of Fundy, are linked in continuum to the local ocean currents. We propose that marine animals utilize all these regions in a manner similar to the Bay of Fundy estuaries and properly designed surveys will reveal their presence. Fish passage studies utilizing the Annapolis estuary low-head, tidal turbine on the Bay of Fundy have shown that turbine related mortality of 20–80% per passage occurs depending on fish species, fish size and the efficiency of turbine operation. We suggest that introduction of tidal turbines into open ocean current systems will cause widespread impact on marine populations resulting in significant declines in abundance.     

Population genetic analysis of white shrimp,Litopenaeus setiferus,using microsatellite genetic markers

The white shrimp (Litopenaeus setiferus) is a commercially and recreationally valuable species, yet little is known of its population structure or genetic diversity. White shrimp are distributed along the Atlantic coast of the United States and from the west coast of Florida to the Bay of Campeche, Mexico. In this study, shrimp were collected from North Carolina, South Carolina (four separate collections were taken from 1995 to 1999), Georgia, the Atlantic and Gulf coasts of Florida, Louisiana, Texas and Mexico. DNA was isolated from these individuals, and genetic variation was assessed at six microsatellite loci. These loci were, for the most part, highly polymorphic with an average expected heterozygosity of 0.68. Deviations from Hardy-Weinberg proportions were observed over all samples, but experimental results suggested the presence of null alleles, which confounded a biological interpretation of this result. Pairwise tests of the similarity of allele frequency distributions and distance measure analyses showed broad-scale genetic homogeneity superimposed over occasional indications of random geographical and temporal differentiation. FST and RST estimates over all loci and samples were 0.002 or less and indicated little population structure. Weak but significant genetic differentiation was evident only between pooled western Atlantic and pooled Gulf of Mexico samples. Within the Gulf of Mexico or within the western Atlantic, the large-scale genetic homogeneity observed may be a consequence of genetic mixing resulting from pelagic larvae and adult migrations, while the random local genetic differentiation may be a result of genetic sampling or experimental sampling error. The weak differentiation between shrimp from the Gulf of Mexico and the western Atlantic can be explained by a relatively recent separation of these two populations and/or small amounts of ongoing gene flow.     

Microsatellite analysis indicates an absence of population structure among Hippoglossus hippoglossus in the north‐west Atlantic

To explore the population structure of Atlantic halibut Hippoglossus hippoglossus , 160 fish from four locations in the north‐west Atlantic (Bay of Fundy, Scotian Shelf, Gulf of St Lawrence and Iceland) were examined for evidence of population structure using 18 microsatellite markers. Pair‐wise F _ST and a model‐based cluster analysis revealed no significant differentiation between samples, although uncertainties surrounding Atlantic halibut reproductive behaviour made it difficult to ascertain that only a single breeding population had been sampled at each location     

Regional differentiation of North American Atlantic salmon at allozyme loci

Allozyme variation was characterised by starch gel electrophoresis at 23 enzyme coding loci and one regulatory locus in Atlantic salmon from 53 rivers in Eastern Canada, encompassing the majority of the species' North American range. Variation among rivers was highly heterogeneous and eight of the 15 polymorphisms showed regionally restricted distributions. Nearest neighbour joining (NJ) analysis and multi‐dimensional scaling suggest six distinct regional groups; Labrador/Ungava, Gulf of Saint Lawrence, Newfoundland (excluding Gulf rivers), the Atlantic shore/Southern Uplands of Nova Scotia, the inner Bay of Fundy, and the outer Bay of Fundy. Approximately 25% of observed genetic variation was distributed among these regions with a weak though significant overall correlation of genetic and geographic distance (Mantel Test, r  = 0·255, P  = 0·005). Collectively, the rivers showed consistent divergence from European populations with strong bootstrap support for the two clusters across loci in the NJ analysis. Mean heterozygosity was 0·061 for both continental groups, but the European population showed more than twice the variation among populations. F _ST values were 0·076 and 0·176 for North America and Europe, respectively, with an overall F _ST of 0·330.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

HARBOR PORPOISE (PHOCOENA PHOCOENA) ABUNDANCE IN ALASKA: BRISTOL BAY TO SOUTHEAST ALASKA, 1991-1993

Between 1991 and 1993, Alaska harbor porpoise ( Phocoena phocoena ) abundance was investigated during aerial surveys throughout much of the coastal and offshore waters from Bristol Bay in the eastern Bering Sea to Dixon Entrance in Southeast Alaska. Line-transect methodology was used, and only those observations made during optimal conditions were analyzed. Survey data indicated densities of 4.48 groups/100 km², or approximately 3,531 harbor porpoises (95% C. I. 2,206-5,651) in Bristol Bay and 0.54 groups/100 km², or 136 harbor porpoises (95% C. I. 11-1,645) for Cook Inlet. Efforts off Kodiak Island resulted in densities of 1.85 groups/100 km², or an abundance estimate of 740 (95% C. I. 259-2,115). Surveys off the south side of the Alaska Peninsula found densities of 2.03 groups/100 km² and an abundance estimate of 551 (95% C. I. 423-719). Surveys of offshore waters from Prince William Sound to Dixon Entrance yielded densities of 4.02 groups/100 km² and an abundance estimate of 3,982 (95% C. I. 2,567-6,177). Combining all years and areas yielded an uncorrected density estimate of 3.82 porpoises per 100 km², resulting in an abundance estimate of 8,940 porpoises (CV = 13.8%) with a 95% confidence interval of 6,746-11,848. Using correction factors from other studies to adjust for animals missed by observers, the total number of Alaska harbor porpoises is probably three times this number.     

Foreword

Allozyme variation was characterised by starch gel electrophoresis at 23 enzyme coding loci and one regulatory locus in Atlantic salmon from 53 rivers in Eastern Canada, encompassing the majority of the species' North American range. Variation among rivers was highly heterogeneous and eight of the 15 polymorphisms showed regionally restricted distributions. Nearest neighbour joining (NJ) analysis and multi‐dimensional scaling suggest six distinct regional groups; Labrador/Ungava, Gulf of Saint Lawrence, Newfoundland (excluding Gulf rivers), the Atlantic shore/Southern Uplands of Nova Scotia, the inner Bay of Fundy, and the outer Bay of Fundy. Approximately 25% of observed genetic variation was distributed among these regions with a weak though significant overall correlation of genetic and geographic distance (Mantel Test, r = 0·255, P = 0·005). Collectively, the rivers showed consistent divergence from European populations with strong bootstrap support for the two clusters across loci in the NJ analysis. Mean heterozygosity was 0·061 for both continental groups, but the European population showed more than twice the variation among populations. F_ST values were 0·076 and 0·176 for North America and Europe, respectively, with an overall F_ST of 0·330.     

VARIABILITY IN THE DIET OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN SCOTTISH WATERS 1992–2003

Interannual, seasonal, and regional variation in the diet of porpoises, Phocoena phocoena , in Scottish (UK) waters was studied using stomach contents of animals stranded between 1992 and 2003. Most samples came from the east coast (including many porpoises killed by bottlenose dolphins), with smaller numbers from the west coast and from Shetland. The most important prey types, in terms of contribution by number and mass, were whiting ( Merlangius merlangus ) and sand eels (Ammodytidae). Multivariate analysis confirmed the existence of regional, seasonal, and interannual variation in diet, as well as differences ( i. e. , biases) related to cause of death. These differences were further explored using univariate analyses. Sand eels were more important in the summer months (quarters 2 and 3) and on the east coast, whereas gadids were more important in winter and in the Shetland area. Some, but not all, observed trends in the numerical importance and size of prey taken were consistent with trends in abundance and size of fish taken during research trawl surveys. There was some evidence that porpoises <1 yr old took more gobies (Gobiidae) and shrimps than did older porpoises. Clupeids (herring Clupea harengus and sprat Sprattus sprattus ) formed a relatively small proportion of the diet, but their importance varied from year to year. Although possible methodological biases prevent firm conclusions, it appears that the importance of clupeids in porpoise diet may have decreased since the 1960s, mirroring the decline in North Sea herring abundance. The recovery of the North Sea herring stock in recent years is not as yet reflected in porpoise diet.     

Reproduction in the common porpoise (Phocoena phocoena) of the North Atlantic

Examination of the reproductive organs from 137 Common porpoises ( Phocoena phocoena ) from the Gulf of St Lawrence, the Bay of Fundy and British waters has enabled observations to be made on reproductive events. Birth in Canadian waters occurs mainly in June and July: mating takes place from june to August . The evidence indicates that females do not become pregnant every year. Birth length is 80 to 90cm; juveniles attain a length of 100 to 110 cm after three to four months. Sexual maturity is reached at a length of about 133 cm in males and 145 cm in females, probably in their third or fourth year. The corpus luteum persists throughout gestation and retrogresses from their histological characteristics. One kind possesses much acellular material and is probably a corpus albicans gravitadionis; the other consists of a mass of obliterated vessels and is the remains either of a corpus luteum of an infertile ovulation or of a lutealized follicle. Evidence indicates that there is seasonal testicular activity, reaching a maximum during July and subsiding in late August.     

Restricted matrilineal gene flow and regional differentiation among Atlantic salmon (Salmo salar L.) populations within the Bay of Fundy,eastern Canada

Matrilineal phylogenetic divergence among Atlantic salmon stocks of the Bay of Fundy in south eastern Canada is investigated. Sequence variation in two regions of the mitochondrial ND1 gene, encompassing 710 base pairs, is described for 168 salmon from 11 rivers. Mean overall haplotype and nucleotide diversity (h and pi) observed are 0.5014 and 0.00095, respectively. Nested clade analysis (NCA) and molecular analysis of variance (AMOVA) both point to highly restricted gene flow among rivers and show the haplotype distribution to be geographically structured. Variation among predefined regions of the Bay (16%) is greater than among populations within these regions (14%) The main regional differentiation occurs between rivers of the geographically isolated inner Minas Basin and those elsewhere in the Bay. Differentiation most probably reflects the pattern and nature of the historical processes associated with post-glacial colonisation of the area by salmon following the last Pleistocene glacial maximum c. 180,00 yrs BP.     

RESPONSE OF NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS) TO PLAYBACK OF CALLS RECORDED FROM SURFACE ACTIVE GROUPS IN BOTH THE NORTH AND SOUTH ATLANTIC

The surface active group (SAG) is the most obvious social interaction of the North Atlantic right whale ( Eubalaena glacialis ). SAGs are typically composed of an adult female with two or more males engaged in social behavior near the surface. Distinct calls, believed to be produced by the female, are associated with these groups. Calls recorded from three North Atlantic right whale SAGs and three South Atlantic right whale ( Eubalaena australis ) SAGs were played back to North Atlantic right whales to determine if these sounds are sufficient to attract males to the groups. Playbacks of gunshot sounds produced by North Atlantic right whales were used as a control stimulus. Thirty-six trials were carried out from 1999 to 2001 in the Bay of Fundy, Canada. Whales approached 27 of 31 SAG playbacks and 0 of 5 gunshot playbacks. Where sex was determined ( n = 28), all approaches to North Atlantic SAG recordings were by males. Individuals ( n = 22) of all age and sex classes approached South Atlantic SAG playbacks. These trials indicate that SAG calls from both populations are sufficient to attract right whales to SAGs and that males and females respond differently to stimuli from the North Atlantic. The difference in response to North and South Atlantic SAG stimuli was unexpected. Novelty, species differences in calls, and different seasonal or behavioral context for the recorded stimuli may be responsible for the differences in response.