Parentage analyses of baleen whales are rare, and although mating systems have been hypothesized for some species, little data on realized male reproductive success are available and the patterns of male reproductive success have remained elusive for most species. Here we combine over 20 years of photo-identification data with high-resolution genetic data for the majority of individual North Atlantic right whales to assess paternity in this endangered species. There was significant skew in male reproductive success compared to what would be expected if mating was random (P < 0.001). The difference was due to an excess of males assigned zero paternities, a deficiency of males assigned one paternity, and an excess of males assigned as fathers for multiple calves. The variance in male reproductive success was high relative to other aquatically mating marine mammals, but was low relative to mammals where the mating system is based on resource- and/or mate-defence polygyny. These results are consistent with previous data suggesting that the right whale mating system represents one of the most intense examples of sperm competition in mammals, but that sperm competition on its own does not allow for the same degree of polygyny as systems where males can control access to resources and/or mates. The age distribution of assigned fathers was significantly biased towards older males (P < 0.05), with males not obtaining their first paternity until approximately 15 years of age, which is almost twice the average age of first fertilization in females (8 years), suggesting that mate competition is preventing younger males from reproducing. The uneven distribution of paternities results in a lower effective population size in this species that already has one of the lowest reported levels of genetic diversity, which may further inhibit reproductive success through mate incompatibility of genetically similar individuals.

The North Atlantic right whale, Eubalaena glacialis (Müller, 1776), is one of the world's most endangered large cetaceans. It is widely believed that Basque whalers caused the most dramatic decline of this species in the western North Atlantic during the early-16th and 17th centuries. Previous osteological analysis of 17 historic bones suggested that 50% of the Basque harvest consisted of right whales and 50% of bowhead whales, Balaena mysticetus L., 1758. This 50:50 ratio has been used to estimate pre-exploitation population size, which has subsequently formed the basis of recovery goals and plans for the North Atlantic right whale. Genetic analysis of 21 bones, 13 identified as right whales and 8 as bowhead whales through osteological examination, indicates that in fact only 1 bone was a right whale and 20 were bowhead whales. Additionally, preliminary microsatellite analyses of this specimen are not consistent with the hypothesis that whaling resulted in the low genetic variation found in this species today. These results differ from what would be expected based on any previous view of Basque whaling, and raise questions regarding the impact of Basque whaling on this species.

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Species biogeography is a result of complex events and factors associated with climate change, ecological interactions, anthropogenic impacts, physical geography, and evolution. To understand the contemporary biogeography of a species, it is necessary to understand its history. Specimens from areas of localized extinction are important, as extirpation of species from these areas may represent the loss of unique adaptations and a distinctive evolutionary trajectory. The walrus (Odobenus rosmarus) has a discontinuous circumpolar distribution in the arctic and subarctic that once included the southeastern Canadian Maritimes region. However, exploitation of the Maritimes population during the 16th-18th centuries led to extirpation, and the species has not inhabited areas south of 55°N for ∼250 years. We examined genetic and morphological characteristics of specimens from the Maritimes, Atlantic (O. r. rosmarus) and Pacific (O. r. divergens) populations to test the hypothesis that the first group was distinctive. Analysis of Atlantic and Maritimes specimens indicated that most skull and mandibular measurements were significantly different between the Maritimes and Atlantic groups and discriminant analysis of principal components confirmed them as distinctive groups, with complete isolation of skull features. The Maritimes walrus appear to have been larger animals, with larger and more robust tusks, skulls and mandibles. The mtDNA control region haplotypes identified in Maritimes specimens were unique to the region and a greater average number of nucleotide differences were found between the regions (Atlantic and Maritimes) than within either group. Levels of diversity (h and π) were lower in the Maritimes, consistent with other studies of species at range margins. Our data suggest that the Maritimes walrus was a morphologically and genetically distinctive group that was on a different evolutionary path from other walrus found in the north Atlantic.

Mitochondrial DNA (mtDNA) sequences were analyzed from 106 bowhead whale (Balaena mysticetus) specimens dating 471 ± 44 14C b.p.–10,290 ± 150 14C b.p. to evaluate whether historical changes in distribution and connectivity were detectable in levels of diversity and population structuring in the Central Canadian Arctic. The species has maintained levels of mtDNA diversity over 10,000 yr comparable to other nonbottlenecked large whale species. When compared to data from the Holocene East Greenland/Spitsbergen and contemporary Bering‐Chuckchi‐Beaufort populations, differentiation was low (FST≤ 0.005, ΦST≤ 0.003) and no temporal or geographical genetic structuring was evident. A combination of analyses suggests that the population has expanded over the past 30,000 14C yr. This genetic signature of expansion could result from population growth, admixture of multiple gene pools, or a combination of both scenarios. Despite known climatic change that altered bowhead distribution and led to isolation of populations, there is no detectable population structuring or change in genetic diversity during the Holocene. This may be due to long generation time, occasional population connectivity and a historically large global population. These characteristics warrant caution when interpreting contemporary bowhead whale DNA data, as it is unlikely that any population will be in mutation‐drift equilibrium.

Low levels of genetic variability identified within the North Atlantic right whale (Eubalaena glacialis), when compared to the Southern right whale (E. australis) and other large whales, have been suggested to result from population reductions due to whaling. Previous genetic analysis of 218 whale bones from sixteenth century Basque whaling sites in the western North Atlantic revealed only a single right whale bone. We determined the genotypes of 27 microsatellite loci using DNA isolated from this bone. All alleles from the historic specimen occur in the extant western North Atlantic population and both the probability of identity of the specimen and the number of heterozygous loci are similar to that in the extant population. Assessments of how genetically different the historical population might have been suggest genetic characteristics have not changed substantially over four centuries of whaling.

Many species of whales went through recent bottlenecks due to commercial whaling. These declines were rapid and recent relative to the life spans and generation times of these species, raising questions regarding to what degree commercial whaling influenced the genetic characteristics of these populations. We analyzed mitochondrial and nuclear DNA from pre‐ and postwhaling samples from two populations that have arguably shown the greatest degree of recovery: eastern North Pacific gray and humpback whales. We also compare the performance of different methods to test for historic bottlenecks and infer past demography based on genetic data. We found substantially higher levels of genetic diversity in gray than in humpback whales (for both time periods), likely due to recent connectivity between Atlantic and Pacific gray whale populations. Other than mitochondrial diversity in humpback whales, levels of diversity were not lower in contemporary samples relative to prewhaling samples, indicating that commercial whaling had a minimal impact on metrics of genetic diversity themselves. However, it did have large impacts on the patterns of diversity, as evidenced by all coalescent‐based methods showing clear evidence of a bottleneck for both populations, whereas all but one method not based on the coalescent failed to detect a bottleneck.

The comment by Romero and Kannada is presented as a critique of our previous work and suggests that our conclusions are in direct conflict with the historic whaling information reviewed in their paper. However, the critique is based on a misinterpretation of the geographical, temporal, and taxonomic focus of our previous work. The source of the putative conflict appears to stem from the misinterpretation that our results, focused solely on the impact of Basque whaling in the 16th and 17th centuries on the western North Atlantic right whale, were intended to be representative of all whaling of both right and bowhead whales throughout the North Atlantic. To demonstrate this, we briefly review our original results and conclusions and show that the information reviewed by Romero and Kannada does not challenge any aspect of our original work. As such, their comment is not a critique of our paper, but rather a brief review of the history of whaling in the North Atlantic.Résumé : Les commentaires de Romero et Kannada se veulent une critique de notre travail antérieur et laissent entendre que nos conclusions sont en contradiction directe avec les données historiques récapitulées dans leur article. Cependant, cette critique se base sur une mauvaise interprétation des échelles géographique, temporelle et taxonomique de notre travail. La source du conflit apparent semble provenir d'une mauvaise impression selon laquelle nos résultats, qui couvrent seulement l'impact de la chasse par les basques sur la baleine franche du Nord de l'ouest de l'Atlantique nord aux 16 e et 17 e siècles, représentent les effets de toute la chasse aux baleines franches du Nord et aux baleines franches boréales dans l'ensemble de l'Atlantique nord. Afin de faire la lumière sur la question, nous passons en revue nos résultats et nos conclusions d'origine et montrons que les données étudiées par Romero et Kannada ne mettent en question aucun aspect de notre travail original. Leurs commentaires ne sont donc pas une critique de notre étude, mais plutôt un bref aperçu de l'histoire de la chasse à la baleine dans l'Atlantique Nord.[Traduit par la Rédaction]