Cetacean mitochondrial DNA control region: sequences of all extant baleen whales and two sperm whale species

The sequence of the mitochondrial control region was determined in all 10 extant species commonly assigned to the suborder Mysticeti (baleen or whalebone whales) and to two odontocete (toothed whale) species (the sperm and the pygmy sperm whale). In the mysticetes, both the length and the sequence of the control region were very similar, with differences occurring primarily in the first approximately 160 bp of the 5' end of the L-strand of the region. There were marked differences between the mysticete and sperm whale sequences and also between the two sperm whales. The control region, less its variable portion, was used in a comparison including the 10 mysticete sequences plus the same region of an Antarctic minke whale specimen and the two sperm whales. The difference between the minke whales from the North Atlantic and the Antarctic was greater than that between any acknowledged species belonging to the same genus (Balaenoptera). The difference was similar to that between the families Balaenopteridae (rorquals) and Eschrichtiidae (gray whales). The findings suggest that the Antarctic minke whale should have a full species status, B. bonaerensis. Parsimony analysis separated the bowhead and the right whale (family Balaenidae) from all remaining mysticetes, including the pygmy right whale. The pygmy right whale is usually included in family Balaenidae. The analysis revealed a close relationship between the gray whale (family Eschrichtiidae) sequence and those of the rorquals (family Balaenopteridae). The gray whale was included in a clade together with the sei, Bryde's, fin, blue, and humpback whales. This clade was separated from the two minke whale types, which branched together.      

Migration of Antarctic minke whales to the Arctic

The Antarctic minke whale (Balaenoptera bonaerensis), and the common minke whale found in the North Atlantic (Balaenoptera acutorostrata acutorostrata), undertake synchronized seasonal migrations to feeding areas at their respective poles during spring, and to the tropics in the autumn where they overwinter. Differences in the timing of seasons between hemispheres prevent these species from mixing. Here, based upon analysis of mitochondrial and microsatellite DNA profiles, we report the observation of a single B. bonaerensis in 1996, and a hybrid with maternal contribution from B. bonaerensis in 2007, in the Arctic Northeast Atlantic. Paternal contribution was not conclusively resolved. This is the first documentation of B. bonaerensis north of the tropics, and, the first documentation of hybridization between minke whale species.     

Morphology and phylogenetic relationships of a new eschrichtiid genus (Cetacea: Mysticeti) from the Early Pliocene of northern Italy

A new eschrichtiid, Eschrichtioides gastaldii gen. nov., comb. nov. , is established based on a specimen previously assigned to Balaenoptera gastaldii Portis, 1885. The holotype is from the Early Pliocene of north-east Italy. It represents a fossil mysticete closely related to the living grey whale, Eschrichtius robustus . Comparative morphology and phylogenetic analysis support the monophyly of Eschrichtiidae and Cetotherium -like mysticetes and a sister group relationship between this clade and Balaenopteridae. Eschrichtiid fossils previously described are all from the Pleistocene and Late Pliocene while Eschrichtioides gastaldii is from the Early Pliocene. The recognition of this new eschrichtiid genus suggests that the Mediterranean trophic web of the Early Pliocene was more complex than at present and that the Neogene mysticete family-level biodiversity of the Mediterranean was higher than that currently observed in this basin.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 161–186.     

A new whale barnacle from the early Pleistocene of Italy suggests an ancient right whale breeding ground in the Mediterranean

The fragmentary fossil history of whale barnacles (Cirripedia: Coronulidae) is mainly constituted by remains of Coronula spp. in Plio-Pleistocene deposits found along extant humpback whale migration routes, and especially in occurrence of breeding areas. Here we report the presence of a single compartment of Cetopirus along with remains of Coronula bifida in Lower Pleistocene open shelf deposits of Salento (South Italy). This is the first occurrence of the genus Cetopirus prior to the Late Glacial period (ca. 15–10 ky before Present), and the specimen here studied is designated as the holotype of the new fossil species Cetopirus fragilis. Since Cetopirus is currently found exclusively inhabiting the skin of the right whales (Eubalaena spp.), the fossil material here studied suggests the persistence of Balaenidae in the Mediterranean Basin during the Early Pleistocene and supports the existence of a baleen whale migratory route active between the central Mediterranean and the North Atlantic during the Plio-Pleistocene.     

Radiation and speciation of pelagic organisms during periods of global warming: the case of the common minke whale, Balaenoptera acutorostrata

How do populations of highly mobile species inhabiting open environments become reproductively isolated and evolve into new species? We test the hypothesis that elevated ocean‐surface temperatures can facilitate allopatry among pelagic populations and thus promote speciation. Oceanographic modelling has shown that increasing surface temperatures cause localization and reduction of upwelling, leading to fragmentation of feeding areas critical to pelagic species. We test our hypothesis by genetic analyses of populations of two closely related baleen whales, the Antarctic minke whale (Balaenoptera bonaerensis) and common minke whale (Balaenoptera acutorostrata) whose current distributions and migration patterns extent are largely determined by areas of consistent upwelling with high primary production. Phylogeographic and population genetic analyses of mitochondrial DNA control‐region nucleotide sequences collected from 467 whales sampled in four different ocean basins were employed to infer the evolutionary relationship among populations of B. acutorostrata by rooting an intraspecific phylogeny with a population of B. bonaerensis. Our findings suggest that the two species diverged in the Southern Hemisphere less than 5 million years ago (Ma). This estimate places the speciation event during a period of extended global warming in the Pliocene. We propose that elevated ocean temperatures in the period facilitated allopatric speciation by disrupting the continuous belt of upwelling maintained by the Antarctic Circumpolar Current. Our analyses revealed that the current populations of B. acutorostrata likely diverged after the Pliocene some 1.5 Ma when global temperatures had decreased and presumably coinciding with the re‐establishment of the polar–equatorial temperature gradient that ultimately drives upwelling. In most population samples, we detected genetic signatures of exponential population expansions, consistent with the notion of increasing carrying capacity after the Pliocene. Our hypothesis that prolonged periods of global warming facilitate speciation in pelagic marine species that depend on upwelling should be tested by comparative analyses in other pelagic species.     

MORPHOMETRIC COMPARISON OF MINKE WHALES BALAENOPTERA ACUTOROSTRATA FROM DIFFERENT AREAS OF THE NORTH ATLANTIC

Multivariate statistical analyses of 18 morphometric characters were performed in order to evaluate potential heterogeneity between predefined minke whale stock unit areas in the North Atlantic (West Greenland, Central, and Northeastern stocks). Results from principal component analyses suggested that the data cannot be regarded as random samples drawn from one uniform distribution. The overlap between groups was too substantial, however, to allow a firm conclusion concerning the question of isolated breeding stocks versus a large common breeding pool.     

A NEW BASAL BALAENOPTERID WHALE FROM THE PLIOCENE OF NORTHERN ITALY

Abstract:  A new basal balaenopterid genus and species, Archaebalaenoptera castriarquati , is described and compared with all the living and fossil members of the family Balaenopteridae and related fossil rorqual-like taxa. It was found in the Lower Pliocene of northern Italy, and is characterized by a supraoccipital with a transversely compressed anterior process, the zygomatic process of the squamosal diverging from the longitudinal axis of the skull, very long nasal bones, and subtle exposition of the parietal on the dorsal wall of the skull. It is primitive in having a maxilla with a long ascending process that is posteriorly unexpanded and round, and a dentary that is straight and not bowed outward, unlike that of living Balaenopteridae. In particular, the discovery of this new genus suggests that, among the early members of Balaenopteridae, the acquisition of the typical sutural pattern shown by maxilla, frontal, parietal and supraoccipital preceded the acquisition of the feeding-related traits that are characteristic of the family. The primitive morphology of the feeding-related structures of A. castriarquati (i.e. the straight dentary and the flat glenoid fossa of the squamosal) suggests that this whale was unable to undertake the intermittent ram feeding typical of Balaenopteridae as efficiently as living members of the family.     

SKULL MORPHOLOGY AND PHYLOGENETIC RELATIONSHIPS OF A NEW DIMINUTIVE BALAENID FROM THE LOWER PLIOCENE OF BELGIUM

Abstract:  A new small balaenid is described and compared to all fossil and living balaenid taxa. The specimen represents a new genus and species and is named Balaenella brachyrhynus . It was discovered in the Lower Pliocene of Kallo (north-west Antwerp, Belgium) and adds new information on the diversity and evolution of Balaenidae. Based on both comparative morphology and phylogenetic analysis, Balaenella brachyrhynus is morphologically closer to the genus Balaena , including the living Greenland Bowhead whale ( B. mysticetus ), and two Pliocene species ( B. montalionis and B. ricei ) from central Italy and the eastern USA. Balaenella brachyrhynus has very short nasals, a short rostrum relative to the total skull length and a horizontal supraoccipital. A cladistic treatment of 81 morphological character states scored for 10 balaenids and nine non-balaenid cetaceans revealed that the other small balaenids generally included in the genus Balaenula (including Balaenula astensis, B. balaenopsis and a Pliocene Balaenula sp. from Japan) are closer to the living genus Eubalaena (the Right whale). As the new skull is so different from the nominal Balaenula species, and as it is more closely related to Balaena than to Eubalaena , it is concluded that a small body size was a common condition in different Balaenidae clades.     

DETERMINATION OF PREGNANCY STATUS FROM BLUBBER SAMPLES IN MINKE WHALES (BALAENOPTERA ACUTOROSTRATA)

Measurement of progesterone concentration in blubber was developed as a method to detect pregnancy in minke whales. Progesterone was extracted and quantified from blubber samples of minke whale carcasses by radioimmunoassay. Results showed a highly significant difference (almost 60-fold) between blubber progesterone concentrations of anatomically determined pregnant females versus non-pregnant female or male carcasses. The results of the study suggest that the blubber progesterone concentrations might be used to determine pregnancy status in free-ranging whales.     

EVOLUTION, TAXONOMY AND ANTITROPICAL DISTRIBUTIONS OF THE PORPOISES (PHOCOENIDAE, MAMMALIA)

True porpoises are a morphologically distinctive and evolutionarily old group of odontocete cetaceans classified as the family Phocoenidae. They are distinct from members of the family Delphinidae, with which they have sometimes been classified. Re-examination of all living and fossil species of phocoenids yields new information on the evolution of the family and indicates the need for significant taxonomic changes. Two separate lines of descent within the group are classified as the subfamilies Phocoeninae and Phocoenoidinae. The living southern hemisphere spectacled porpoise, Phocoena dioptrica , is actually more closely related to the North Pacific Dali's porpoise, Phocoenoides dalli , than to other species of Phocoena. It therefore belongs in the subfamily Phocoenoidinae with Phocoenoides but represents a separate genus, Australophocaena (new genus). The earliest unquestioned fossil phocoenids have been found only in rocks around the Pacific basin. They include latest Miocene and Pliocene species of Piscolithax and the Late Miocene Salumiphocaena stocktoni (new genus) in the Phocoenoidinae, and an undescribed latest Miocene species in the Phocoeninae. World climate has influenced past and present distributions of phocoenids, and Australopbocaena dioptrica is possibly the antitropical counterpart of Phocoenoides dalli. All species of living phocoenids show similarities caused by convergent evolution, resulting in part from the phenomenon of paedomorphosis.     

Exploring the Formation of a Disjunctive Pattern between Eastern Asia and North America Based on Fossil Evidence from Thuja (Cupressaceae)

Thuja, a genus of Cupressaceae comprising five extant species, presently occurs in both East Asia (3 species) and North America (2 species) and has a long fossil record from Paleocene to Pleistocene in the Northern Hemisphere. Two distinct hypotheses have been proposed to account for the origin and present distribution of this genus. Here we recognize and describe T. sutchuenensis Franch., a new fossil Thuja from the late Pliocene sediments of Zhangcun, Shanxi, North China, based on detailed comparisons with all living species and other fossil ones, integrate the global fossil records of this genus plotted in a set of paleomaps from different time intervals, which show that Thuja probably first appeared at high latitudes of North America in or before the Paleocene. This genus reached Greenland in the Paleocene, then arrived in eastern Asia in the Miocene via the land connection between East Asia and western North America. In the late Pliocene, it migrated into the interior of China. With the Quaternary cooling and drying, Thuja gradually retreated southwards to form today’s disjunctive distribution between East Asia and North America.     

IN VITRO FERTILIZATION OF IN VITRO MATURED MINKE WHALE (BALAENOPTERA ACUTOROSTRATA) FOLLICULAR OOCYTES

In vitro fertilization of follicular oocytes harvested from ovaries and matured in vitro was attempted for 55 minke whales ( Balaenoptera acutorostrata ) captured for Japanese research purposes in the Antarctic Ocean during the period from November 1995 to March 1996. In Experiment 1, effects of culture duration (96 h or 120 h) on maturation of follicular oocytes and addition of caffeine (5 mM) and/or heparin (100 pg/ml) on sperm penetration and pro-nuclear formation were investigated. Spermatozoa recovered from the vasa deferentia of four mature males were diluted (5-fold) and frozen at - 80°C. The post-thawed and pooled spermatozoa were used for in vitro insemination. A higher ( P < 0.05) proportion of the oocytes cultured for 120 h (34.2% of 260) progressed beyond the second metaphase stage than of the oocytes cultured for 96 h (26.0% of 262). For the matured oocytes, higher rates of penetration ( P < 0.05) and pronuclear formation ( P < 0.01) were obtained in the oocytes cultured for 120 h (55.1% and 40.4%) than in those cultured for 96 h (32.4 % and 20.6%). Addition of caffeine and heparin did not show a significant effect. In Experiment 2, follicular oocytes matured for 120 h and then inseminated were cultured to examine the subsequent development in two culture systems (with and without co-cultured cumulus cells). Of 448 inseminated oocytes, cleaved embryos (2–16 cells) were observed with (5.8%) and without (4.9%) co-cultured systems. No cleavage was observed in 54 ova without insemination. These results indicate that in vitro fertilization of minke whale in vitro matured follicular oocytes with cryopreserved spermatozoa is possible, yielding cleaved embryos.     

HEMATOLOGY AND BLOOD CHEMISTRY OF A BRYDE'S WHALE, BALAENOPTERA EDENI, ENTRAPPED IN THE MANNING RIVER, NEW SOUTH WALES, AUSTRALIA

A Bryde's whale, Balaenoptera edeni , was rescued after having been entrapped in the Manning River, Australia, for 100 d. Blood, skin, and epidermal tissue were analyzed to determine the whale's taxonomic status, gender, and health. This paper reports the results of these analyses and discusses the findings in relation to the potential physiological damage to the whale from its protracted stay in the river. Molecular genetic analysis of epidermal tissue identified the whale as a male of the rare pygmy form of Bryde's whale. The 10.3-m whale was diagnosed as emaciated, parasitized, under stress, and in a state of profound catabolism at the time of its rescue. Although it had greatly depleted its energy reserves, the whale was rescued from the river before the onset of ill health or irreparable physiological damage. No organ dysfunction was evident with the possible exception of some deterioration in kidney function most likely caused by parasitism. The whale swam away srongly on release, and its chances of survival appeared to be good.     

Cetacean relaxin. Isolation and sequence of relaxins from Balaenoptera acutorostrata and Balaenoptera edeni

The tendency toward extremely high variability among relaxins derived from purportedly closely related species has come to an abrupt end with the discovery of quasi-porcine relaxin in the minke whale (Balaenoptera acutorostrata) and the Bryde's whale (Balaenoptera edeni). An aqueous abstract of the corpora lutea of the two baleen whales contained significant amounts of relaxin-like activity as determined by a mouse bioassay and by cross-reactivity with anti-pig relaxin antibodies. The activity could be isolated and purified to homogeneity. Sequence analysis revealed that both whale relaxins differed from each other by about 3 residues, whereas the relaxin of B. edeni differed at only one position from that of pig relaxin. The similarity appears to include even the chain length heterogeneity observed at the C-terminal end of the B chain in porcine relaxin which is produced by a peculiar mode of connecting peptide removal from the pro-hormone. This finding may well represent one of the better documented challenges to the current paradigm of molecular evolution.     

Population structure and possible migratory links of common minke whales, Balaenoptera acutorostrata, in the Southern Hemisphere

Mitochondrial DNA control region sequences were analyzed to investigate population structure and possible migratory links of common minke whales (Balaenoptera acutorostrata) in two ocean basins: western South Atlantic (WSA) and western South Pacific (WSP). The results of several different phylogenetic estimations consistently grouped all haplotypes but one (n = 1) from these two ocean basins into two separate clades. South and North Atlantic haplotypes were more closely related to each other than either was to haplotypes from the WSP. The interpopulation genetic distance between WSA and WSP whales was similar to that reported between North Pacific and North Atlantic common minke whales (0.0234). The migration rate between the two ocean basins was estimated at near-zero using MDIV. The genetic evidence presented here was consistent with the hypothesis of migratory links among Brazil, Chilean Patagonia and the Antarctic Peninsula, and between low-latitude and Antarctic waters of the WSP. The results suggest multiple populations of common minke whales in the Southern Hemisphere, which may have conservation as well as taxonomic implications. Our single locus results should be corroborated by additional analyses in a larger number of samples and at more genetic markers.     

Analysis of the C4 genes in baleen whales using a human cDNA probe

We have used a human C4 cDNA probe to investigate the complement component C4 gene in four members of the family Balaenopteridae: fin whale (Balaenoptera physalus), sei whale (B. borealis), minke whale (B. acutorostrata), and bryde's whale (B. edeni). Restriction mapping of genomic DNA from the first three species suggests the presence of only one locus in these species, and also shows that the C4 genes in the three species are very similar. We have used 14 restriction endonucleases to investigate the restriction fragment length polymorphism (RFLP) of fin whales, 13 enzymes for sei whales, and 8 enzymes for the minke whale. No polymorphism was seen in DNA from the five minke whale samples, but Rsa I and Taq I restriction enzymes gave polymorphism in fin and sei whales whereas Hind III and Msp I restriction enzymes showed polymorphism in sei whales only. Only one bryde's whale sample was available for investigation. The study of DNA available from mother-fetus pairs from the two polymorphic species demonstrated a simple, two-allele transmission of RFLP alleles.     

Chimeric structure of omp2 of Brucella from Pacific common minke whales (Balaenoptera acutorostrata)

In the Pacific common minke whale (Balaenoptera acutorostrata ), a new variant of Brucella has been detected using the polymerase chain reaction. Detailed analysis of the porin protein genes, omp2a and omp2b from the whale Brucella showed that these two genes have some motifs in common with Atlantic marine strains in the 5'-terminal one-third region. On the other hand, the nucleotide sequences in the 3'-terminal two-thirds region of the two genes were almost identical to the respective genes of terrestrial strains. Thus, Pacific whale Brucella omp2 genes are chimeras between marine and terrestrial strains.     

The cardiac ventricles of a baleen whale (Balaenoptera acutorostrata: Minke whale) and a toothed whale (Hyperoodon ampullatus: Bottlenose whale)

Fifteen minke whale (Balaenoptera acutorostrata: Mysticeti) and three bottlenose whale (Hyperoodon ampullatus: Odontoceti) hearts were obtained from Norway. Only the ventricular mass and the base of the great arteries were available for study. The shape of the heart, the double apex, the narrowness of the interventricular septum in the dorsiventral plane, the side-to-side position of the ventricles, the relative thickness of the wall of the right ventricle, and the heavy trabeculation agree with other, brief reports of whale heart morphology. Coronary artery tortuosity, which has been described in sperm whales by others, was found in the bottlenose but not in the minke whales. It is suggested that the shape of the heart is associated with depth and duration of dives and that the pattern of trabeculation may be related to the size of the animal.     

Rorqual whale (Balaenopteridae) surface lunge‐feeding behaviors: Standardized classification,repertoire diversity,and evolutionary analyses

Rorqual whales (Family: Balaenopteridae) are the world's largest predators and sometimes feed near or at the sea surface on small schooling prey. Most rorquals capture prey using a behavioral process known as lunge‐feeding that, when occurring at the surface, often exposes the mouth and head above the water. New technology has recently improved historical misconceptions about the natural variation in rorqual lunge‐feeding behavior yet missing from the literature is a dedicated study of the identification, use, and evolution of these behaviors when used to capture prey at the surface. Here we present results from a long‐term investigation of three rorqual whale species (minke whale, Balaenoptera acutorostrata; fin whale, B. physalus; and blue whale, B. musculus) that helped us develop a standardized classification system of surface lunge‐feeding (SLF) behaviors. We then tested for differences in frequency of these behaviors among the three species and across all rorqual species. Our results: (1) propose a unified classification system of six homologous SLF behaviors used by all living rorqual whale species; (2) demonstrate statistically significant differences in the frequency of each behavior by minke, fin, and blue whales; and (3) provide new information regarding the evolution of lunge‐feeding behaviors among rorqual whales.