The effect of differential reproductive success on population genetic structure: correlations of life history with matrilines in humpback whales of the gulf of maine

To examine whether demographic and life-history traits are correlated with genetic structure, we contrasted mtDNA lineages of individual humpback whales (Megaptera novaeangliae) with sighting and reproductive histories of female humpback whales between 1979 and 1995. Maternal lineage haplotypes were obtained for 323 whales, either from direct sequencing of the mtDNA control region (n = 159) or inferred from known relationships along matrilines from the sequenced sample of individuals (n = 164). Sequence variation in the 550 bp of the control region defined a total of 19 maternal lineage haplotypes that formed two main clades. Fecundity increased significantly over the study period among females of several lineages among the two clades. Individual maternal lineages and other clades were characterized by significant variation in fecundity. The detected heterogeneity of reproductive success has the potential to substantially affect the frequency and distribution of maternal lineages found in this population over time. There were significant yearly effects on adult resighting rates and calf survivorship based on examination of sighting histories with varying capture-recapture probability models. These results indicate that population structure can be influenced by interactions or associations between reproductive success, genetic structure, and environmental factors in a natural population of long-lived mammals.     

Preliminary analysis of mitochondrial DNA variation in a southern feeding group of eastern North Pacific gray whales

Although the majority of eastern North Pacific(ENP) gray whales migrate to feeding grounds inthe Bering and Chukchi Seas, some terminate themigration in more southerly areas such asBritish Columbia (BC). Long-term sightingstudies in Clayoquot Sound (CS), BC, indicatethat approximately 35–50 individuals exhibitlong-term fidelity to this site. To determinethe sex composition (based on genetic sexing)of CS gray whales and to assess whethermatrilineal site-fidelity occurs in CS, wecollected skin biopsy samples from 16 CSindividuals (`residents') and 41 samples fromother areas (representative of the overallpopulation in the ENP: `non-residents'). Atotal of 27 polymorphic sites defined 24haploytpes among the 57 samples sequenced forHV1 of the mtDNA control region. Thenucleotide and haplotype diversities of thesesamples were 0.017 (SE = 0.0012) and 0.94(SE = 0.0019), respectively. Neighbor-joininganalysis revealed five lineages each of whichcontained haplotypes that were observed in bothresidents and non-residents. Residents did notdiffer significantly from non-residents, and nosignificant sex-ratio bias was found. Thesedata suggest a level of diversity that isinconsistent with a severe historicalbottleneck, and given the available samplesize, do not indicate matrilineally directedfidelity to Clayoquot Sound.     

African-American mitochondrial DNAs often match mtDNAs found in multiple African ethnic groups

Background  

Mitochondrial DNA (mtDNA) haplotypes have become popular tools for tracing maternal ancestry, and several companies offer this service to the general public. Numerous studies have demonstrated that human mtDNA haplotypes can be used with confidence to identify the continent where the haplotype originated. Ideally, mtDNA haplotypes could also be used to identify a particular country or ethnic group from which the maternal ancestor emanated. However, the geographic distribution of mtDNA haplotypes is greatly influenced by the movement of both individuals and population groups. Consequently, common mtDNA haplotypes are shared among multiple ethnic groups. We have studied the distribution of mtDNA haplotypes among West African ethnic groups to determine how often mtDNA haplotypes can be used to reconnect Americans of African descent to a country or ethnic group of a maternal African ancestor. The nucleotide sequence of the mtDNA hypervariable segment I (HVS-I) usually provides sufficient information to assign a particular mtDNA to the proper haplogroup, and it contains most of the variation that is available to distinguish a particular mtDNA haplotype from closely related haplotypes. In this study, samples of general African-American and specific Gullah/Geechee HVS-I haplotypes were compared with two databases of HVS-I haplotypes from sub-Saharan Africa, and the incidence of perfect matches recorded for each sample.     

Hierarchical structure of mitochondrial DNA gene flow among humpback whales Megaptera novaeangliae, world-wide

The genetic structure of humpback whale populations and subpopulation divisions is described by restriction fragment length analysis of the mitochondrial (mt) DNA from samples of 230 whales collected by biopsy darting in 11 seasonal habitats representing six subpopulations, or 'stocks', world-wide. The hierarchical structure of mtDNA haplotype diversity among population subdivisions is described using the analysis of molecular variance (AMOVA) procedure, the analysis of gene identity, and the genealogical relationship of haplotypes as constructed by parsimony analysis and distance clustering. These analyses revealed: (i) significant partitioning of world-wide genetic variation among oceanic populations, among subpopulations or 'stocks' within oceanic populations and among seasonal habitats within stocks; (ii) fixed categorical segregation of haplotypes on the south-eastern Alaska and central California feeding grounds of the North Pacific; (iii) support for the division of the North Pacific population into a central stock which feeds in Alaska and winters in Hawaii, and an eastern or 'American' stock which feeds along the coast of California and winters near Mexico; (iv) evidence of genetic heterogeneity within the Gulf of Maine feeding grounds and among the sampled feeding and breeding grounds of the western North Atlantic; and (v) support for the historical division between the Group IV (Western Australia) and Group V (eastern Australia, New Zealand and Tonga) stocks in the Southern Oceans. Overall, our results demonstrate a striking degree of genetic structure both within and between oceanic populations of humpback whales, despite the nearly unlimited migratory potential of this species. We suggest that the humpback whale is a suitable demographic and genetic model for the management of less tractable species of baleen whales and for the general study of gene flow among long-lived, mobile vertebrates in the marine ecosystem.     

Mitochondrial DNA diversity and population structure among southern right whales (Eubalaena australis)

The population structure and mitochondrial (mt) DNA diversity of southern right whales (Eubalaena australis) are described from 146 individuals sampled on 4 winter calving grounds (Argentina, South Africa, Western Australia, and the New Zealand sub-Antarctic) and 2 summer feeding grounds (South Georgia and south of Western Australia). Based on a consensus region of 275 base pairs of the mtDNA control region, 37 variable sites defined 37 unique haplotypes, of which only one was shared between regional samples of the Indo-Pacific and South Atlantic Oceans. Phylogenetic reconstruction of the southern right whale haplotypes revealed 2 distinct clades that differed significantly in frequencies between oceans. An analysis of molecular variance confirmed significant overall differentiation among the 4 calving grounds at both the haplotype and the nucleotype levels (F(ST) = 0.159; Phi(ST) = 0.238; P < 0.001). Haplotype diversity was significantly lower in the Indo-Pacific (h = 0.701 +/- 0.037) compared with the South Atlantic (h = 0.948 +/- 0.013), despite a longer history of exploitation and larger catches in the South Atlantic. In fact, the haplotype diversity in the Indo-Pacific basin was similar to that of the North Atlantic right whale that currently numbers about 300 animals. Multidimensional scaling of genetic differentiation suggests that gene flow occurred primarily between adjacent calving grounds within an ocean basin, with mixing of lineages from different calving grounds occurring on feeding grounds.     

Population spatial structuring on the feeding grounds in North Atlantic humpback whales (Megaptera novaeangliae)

Population spatial structuring among North Atlantic humpback whales Megaptera novaeangliae on the summer feeding grounds was investigated using movement patterns of identified individuals. We analysed the results from an intensive 2-year ocean-basin-scale investigation resulting in 1658 individuals identified by natural markings and 751 individuals by genetic markers supplemented with data from a long-term collaborative study with 3063 individuals identified by natural markings. Re-sighting distances ranged from <1 km to >2200 km. The frequencies ( F ) of re-sighting distances ( D ) observed in consecutive years were best modelled by an inverse allometric function ( F =6631 D ^−1.24, r ²=0.984), reflecting high levels of site fidelity (median re-sighting distance <40 km) with occasional long-distance movement (5% of re-sightings >550 km). The distribution of re-sighting distances differed east and west of 45°W, with more long-distance movement in the east. This difference is consistent with regional patterns of prey distribution and predictability. Four feeding aggregations were identified: the Gulf of Maine, eastern Canada, West Greenland and the eastern North Atlantic. There was an exchange rate of 0.98% between the western feeding aggregations. The prevalence of long-distance movement in the east made delineation of possible additional feeding aggregations less clear. Limited exchange between sites separated by as little as tens of kilometres produced lower-level structuring within all feeding aggregations. Regional and temporal differences in movement patterns reflected similar foraging responses to varying patterns of prey availability and predictability. A negative relationship was shown between relative abundance of herring and sand lance in the Gulf of Maine and humpback whale movement from the Gulf of Maine to eastern Canada.     

Mitochondrial DNA diversity in North American and European Atlantic salmon with emphasis on the Downeast rivers of Maine

The displacement loop and NADH-1 dehydrogenase regions of mitochondrial DNA (mtDNA) were amplified by the polymerase chain reaction in 954 Atlantic salmon and digested with 40 restriction endonucleases. Variation was detected with 10 enzymes, resulting in 21 composite haplotypes which were strongly patterned geographically with a major discontinuity observed between most North American (NA) and European salmon. Significant heterogeneity of haplotype frequencies was found within and among all classification levels (continent, country, and river). Haplotype frequencies were significantly different across continents, within European samples, within NA samples, within Canadian samples, within wild Maine samples, within captive Maine strains, and between captive and wild Maine strains. Nine haplotypes occurred only in NA, seven in Maine, three only in Maine, and 11 occurred only in Europe. Some Maine rivers had only a single haplotype, suggesting that effective population sizes may be low. The second most frequent European haplotype occurred in tributaries to one Newfoundland river. Gene trees based on parsimony and genetic distance suggest that the haplotypes are monophyletic within each continent, and that the haplotype found on both continents is intermediate between those of Europe and NA, suggesting common ancestry of all haplotypes.     

Diversity in and evidence for selection on the mitochondrial genome of Phytophthora infestans

Two extant nomenclature systems were reconciled to relate six mitochondrial DNA (mtDNA) haplotypes of Phytophthora infestans, the oomycete pathogen causing late blight disease on potato and tomato. Carter's haplotypes I-a and I-b were included in Goodwin's haplotype A, while Carter's haplotypes II-a and II-b were included in Goodwin's haplotype B. In addition, haplotypes E and F were included in Carter's haplotype I-b. The mutational differences separating the various haplotypes were determined, and we propose that either haplotype I-b(A) or haplotype I-a(A) is the putative ancestral mtDNA of P. infestans, because either can center all the other haplotypes in a logical stepwise network of mutational changes. The occurrence of the six haplotypes in 548 isolates worldwide was determined. Haplotypes I-a and II-a were associated with diverse genotypes worldwide. As previously suggested, haplotype I-b was found only in the US-1 clonal lineage and its variants (n = 99 isolates from 16 countries on 5 continents), and haplotype II-b was limited to the US-6 clonal lineage and its derivatives (n = 36). In a confirmation of a previous suggestion, the randomly mating population in the Toluca Valley of central Mexico (n = 78) was monomorphic for mtDNA haplotype I-a(A). We hypothesize that selection there may be driving the dominance of that single mtDNA haplotype.     

Simple identification of mitochondrial lineages in contact zones based on lineage-selective primers

A variety of research projects focus on genetic variation among and within maternal lineages as encompassed by mitochondrial DNA (mtDNA). While mtDNA often differs substantially between species, large differences may also be found within species. The evaluation of such divergent lineages, for example in intraspecific contact zones (hybrid zones), commonly involves sequencing numerous individuals. Large‐scale sequencing is both expensive and labour‐intensive. Based on sequences from 15 individuals, we devised a simple and quick polymerase chain reaction assay for identification of divergent mtDNA lineages in a secondary contact zone of the side‐blotched lizard (Uta stansburiana). The application uses lineage‐selective primers to amplify a lineage‐diagnostic product, and is based on each group of mtDNA haplotypes being a monophyletic assemblage of haplotypes sharing the same maternal ancestry, deeply divergent from the other group. The assay was tested on a larger sample (n = 147) of specimens from the contact zone, confirming its usefulness in quick and reliable identification of mtDNA lineages. This approach can be modified for other species, provided diagnostic lineage variation is available, and may also be performed in simple laboratory settings while conducting fieldwork.     

Circumpolar diversity and geographic differentiation of mtDNA in the critically endangered Antarctic blue whale (Balaenoptera musculus intermedia)

The Antarctic blue whale (Balaenoptera musculus intermedia) was hunted to near extinction between 1904 and 1972, declining from an estimated initial abundance of more than 250,000 to fewer than 400. Here, we describe mtDNA control region diversity and geographic differentiation in the surviving population of the Antarctic blue whale, using 218 biopsy samples collected under the auspices of the International Whaling Commission (IWC) during research cruises from 1990–2009. Microsatellite genotypes and mtDNA sequences identified 166 individuals among the 218 samples and documented movement of a small number of individuals, including a female that traveled at least 6,650 km or 131° longitude over four years. mtDNA sequences from the 166 individuals were aligned with published sequences from 17 additional individuals, resolving 52 unique haplotypes from a consensus length of 410 bp. From this minimum census, a rarefaction analysis predicted that only 72 haplotypes (95% CL, 64, 86) have survived in the contemporary population of Antarctic blue whales. However, haplotype diversity was relatively high (0.968±0.004), perhaps as a result of the longevity of blue whales and the relatively recent timing of the bottleneck. Despite the potential for circumpolar dispersal, we found significant differentiation in mtDNA diversity (F_ST = 0.032, p<0.005) and microsatellite alleles (F_ST = 0.005, p<0.05) among the six Antarctic Areas historically used by the IWC for management of blue whales.     

Male-only care and classical polyandry in birds: phylogeny, ecology and sex differences in remating opportunities

It has been argued recently that the combination of male-only parental care and classical polyandry in birds is the most interesting and yet the least understood of all avian breeding systems. Despite a huge number of hypotheses, careful comparative analyses have repeatedly failed to identify consistent ecological differences between species showing male-only care and closely related species showing other patterns of care. This has led to the suggestion that such analyses fail because the crucial differences are between ancient lineages rather than between closely related species. Here, therefore, I use comparisons between families to test three well-known hypotheses: that male-only care is associated with: (i) a low rate of fecundity; (ii) large egg size relative to female size; or (iii) female-biased opportunities for remating. Families showing male-only care do not differ from families showing female-only care with respect to rate of fecundity or relative egg size. There is, however, a significant difference between these two groups of families with respect to an index of remating opportunities, nesting density. Families showing female-only care nest at high density, while those showing male-only care nest at very low density. This is one of the first times a consistent ecological correlate has been identified for male-only care in birds. It suggests that female-only care arises (or persists) in families where remating opportunities are abundant for both sexes, whereas male-only care arises (or persists) in families where remating opportunities are rare for both sexes and particularly scarce for males. This in turn suggests that sex differences in remating opportunities are the key ecological factor in determining male-only care and classical polyandry in birds.     

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.     

mtDNA variation of the critically endangered hawksbill turtle (Eretmochelys imbricata) nesting on Iranian islands of the Persian Gulf

Genetic diversity of sea turtles (hawksbill turtle) was studied using sequencing of mitochondrial DNA (mtDNA, D-loop region). Thirty dead embryos were collected from the Kish and Qeshm Islands in the Persian Gulf. Analysis of sequence variation over 890 bp of the mtDNA control region revealed five haplotypes among 30 individuals. This is the first time that Iranian haplotypes have been recorded. Nucleotide and haplotype diversity was 0.77 and 0.001 for Qeshm Island and 0.64 and 0.002 for Kish Island, respectively. Total haplotype diversity was calculated as 0.69, which demonstrates low genetic diversity in this area. The data also indicated very high rates of migration between the populations of these two islands. A comparison of our data with data from previous studies downloaded from a gene bank showed that turtles of the Persian Gulf migrated from the Pacific and the Sea of Oman into this area. On the other hand, evidence of migration from populations to the West was not found.     

An application of control region sequence as a matrilineage marker for Elliot's pheasant of a zoo population

Control region sequence, an mtDNA marker, was usually used in phylogenesis analysis in species level or genetic structure study among populations. In this study, enlightened by its character of maternal heredity in vertebrates, we used control region sequence as a matrilineage marker for Elliot's pheasant (Syrmaticus ellioti) of Ningbo Zoo population. In Ningbo Zoo, 36 individuals of Elliot's pheasant were descendants from three female founders introduced in 1988. Three control region haplotypes (Ha, Hb, Hc) were identified by six variable nucleotide positions among the control region sequences over 36 individuals. The number of haplotypes was accorded with the number of female founders. Total 20 individuals (C04, C06, C08-11, C14, C20, C21, C23-29, C32, C34-36) shared haplotype a, while 12 individuals (C01, C05, C07, C12, C13, C16-19, C22, C30, C33) shared haplotype b and 4 individuals (C02, C03, C15, C31) shared haplotype c. Those individuals sharing the same haplotype were offspring from one female founder. In other words, there were three maternal lineages and the simple relationship among individuals was indicated. As a result, it seemed that the control region sequence was a useful marker for identification of matrilineage in this study. Meanwhile, the matrilineage information may be compensatory data if there were no any pedigree records in captive species for breeding management.     

Vessel collision injuries on live humpback whales,Megaptera novaeangliae,in the southern Gulf of Maine

North Atlantic humpback whales (Megaptera novaeangliae) in the Gulf of Maine overlap with both recreational and commercial vessel activity. Vessel strikes are one source of anthropogenic impact that has the potential to inhibit the recovery of this protected species. There are currently no regulations or guidelines specifically devised to reduce the likelihood of collisions for vessels transiting in the vicinity of humpback whales, except for vessels actively engaged in whale watching. To understand interactions between vessels and humpback whales better, we analyzed injuries on 624 individuals photographed in the southern Gulf of Maine from 2004 to 2013. Multiple reviewers evaluated 210,733 photos for five categories of injury consistent with a vessel strike. In total, 14.7% (n = 92) of individuals photographed showed injuries consistent with one or more vessel strikes. These results likely underestimate vessel collision rates and impacts because multiple events, events resulting in mortality, and those that involved only blunt force trauma could not be detected. Nevertheless, our results indicate that vessel strikes are underreported and that healing is dependent on the severity and location of the injury. We recommend that a management strategy be developed for all classes of vessels transiting in the vicinity of whales.     

Assessment of genetic structure among eastern North Pacific gray whales on their feeding grounds

Although most eastern North Pacific (ENP) gray whales feed in the Bering, Beaufort, and Chukchi Seas during summer and fall, a small number of individuals, referred to as the Pacific Coast Feeding Group (PCFG), show intra‐ and interseasonal fidelity to feeding areas from northern California through southeastern Alaska. We used both mitochondrial DNA (mtDNA) and 12 microsatellite markers to assess whether stock structure exists among feeding grounds used by ENP gray whales. Significant mtDNA differentiation was found when samples representing the PCFG (n = 71) were compared with samples (n = 103) collected from animals feeding further north (F_ST = 0.012, P = 0.0045). No significant nuclear differences were detected. These results indicate that matrilineal fidelity plays a role in creating structure among feeding grounds but suggests that individuals from different feeding areas may interbreed. Haplotype diversities were similar between strata (h_PCFG = 0.945, h_Northern = 0.952), which, in combination with the low level of mtDNA differentiation identified, suggested that some immigration into the PCFG could be occurring. These results are important in evaluating the management of ENP gray whales, especially in light of the Makah Tribe's proposal to resume whaling in an area of the Washington coast utilized by both PCFG and migrating whales.     

Mitochondrial sequence variation in the Guahibo Amerindian population from Venezuela

New data were obtained on mitochondrial DNA (mtDNA) from Guahibo from Venezuela, a group so far not studied using molecular data. A population sample (n = 59) was analyzed for mtDNA variation in two control-region hypervariable segments (HV1 and HV2) by sequencing. The presence or absence of a 9-bp polymorphism in the COII/tRNA(Lys) region was studied by direct amplification and electrophoretic identification. Thirty-eight variable sites were detected in regions HV1 and HV2, defining 26 mtDNA lineages; 23.7% of these were present in a single individual. The 9-bp deletion was found in 3.39% of individuals. Nucleotide and haplotype diversities were relatively high compared with other New World populations. The identified sequence haplotypes were classified into four major haplogroups (A-D) according to previous studies, with high frequencies for A (47.46%) and C (49.15%), low frequency for B (3.39%), and an absence of D.     

Consequences of a catadromous life-strategy for levels of mitochondrial DNA differentiation among populations of the Australian bass, Macquaria novemaculeata

The influence of a catadromous life-strategy on levels of spatial genetic structuring in fish is poorly understood. In an effort to gain a better appreciation of how this specialized life-strategy determines population genetic structuring, we assessed variation in the mitochondrial DNA (mtDNA) control region in a catadromous perciform, the Australian bass Macquaria novemaculeata . Nineteen putative haplotypes were resolved using temperature gradient gel electrophoresis from 10 geographically distinct populations. Significant heterogeneity was revealed in haplotype frequencies and their spatial distributions among many locales. Gene partitioning statistics ( AMOVA ) for both raw haplotype frequency data and frequency data with sequence divergences were concordant, indicating that M. novemaculeata populations were moderately genetically structured (Φ_ST = 0.05, 0.06; P < 0.001, respectively). Isolation by distance seems to be a strong structuring force in M. novemaculeata , culminating in no detectable phylogeographic structuring among haplotypes. Low sequence divergences were observed among many haplotypes and it is suggested that these are the result of pruning of maternal lineages by cyclical variations in female reproductive success. This study highlights the importance of life-history patterns and, in particular, spawning locality, in determining spatial structuring of mtDNA variation in catadromous species.     

Genetic diversity and population structure of wintering Western Sandpipers from the Sinaloa coast,Mexico

ABSTRACT Although Western Sandpipers (Calidris mauri) are one of the best‐studied shorebirds along the Pacific Flyway, their genetic identity and population structure is poorly known. We studied the genetic population structure of nonbreeding Western Sandpipers at Bahia Santa María in Sinaloa, northwestern Mexico. A 685‐bp fragment of the mtDNA control region was sequenced for 162 individuals, resulting in 29 variable positions that defined 41 haplotypes. The most common haplotype (WESA01) occurred in 44% of all individuals, whereas 15 were unique to single individuals. Nucleotide diversity was low (π= 0.0030 ± 0.0019 [SE]), but haplotype diversity was moderately high (h= 0.802 ± 0.033 [SE]). The main two maternal lineages exhibited a slightly different local scale distribution that appeared to be related to migratory chronology. Lineage A represented 71% of the haplotypes and was evenly distributed across the nonbreeding season and habitat types, whereas Lineage B represented only 29% of the haplotypes and was disproportionately represented in January in some habitat types. Overall, the low level of nucleotide diversity, the star‐shape of the haplotype tree, the mismatch distribution, and the significantly negative Tajima's D values suggest that Western Sandpipers underwent a recent demographic expansion. Although our results are based on a small sample size from one of several wintering sites along the Pacific Flyway, Santa María likely contains a relatively high proportion of the species genetic variability because it is the wintering ground of nearly 10% of the global population.