Comparable length at weaning in cetaceans

Weaning represents the transition of mammalian life from maternal dependence to independence in both energetics and behavior. The length at weaning (Lw) is determined by the maternal investment during gestation and lactation. It affects calf survival and impacts on the long‐term persistence of species, but the measurement is not readily obtainable for many cetaceans. A general linear model and residual correlation were used to explore the correlations between the published Lw and five other life history traits of cetaceans, including female asymptotic length (Lx), length at birth (Lb), lactation period (LP), gestation period (GP), and calving interval (CI). Lx was a significant confounding parameter on the other five traits. By eliminating the confounding Lx, ANCOVA revealed that baleen whales (Mysticeti) and toothed cetaceans (Odontoceti) had comparable Lw. By residual correlation, which factors the confounding Lx, Lw was significantly correlated with Lb but not with GP, LP, and CI. After Lb was further eliminated, convergent Lw in cetaceans could still be observed. Therefore, we proposed a generic expression Lw= 1.239Lx^0.877, which allows us to further estimate Lw of undocumented cetacean species.     

Predicting Cetacean Habitats from Their Energetic Needs and the Distribution of Their Prey in Two Contrasted Tropical Regions

To date, most habitat models of cetaceans have relied on static and oceanographic covariates, and very few have related cetaceans directly to the distribution of their prey, as a result of the limited availability of prey data. By simulating the distribution of six functional micronekton groups between the surface and ≃1,000 m deep, the SEAPODYM model provides valuable insights into prey distributions. We used SEAPODYM outputs to investigate the habitat of three cetacean guilds with increasing energy requirements: sperm and beaked whales, Globicephalinae and Delphininae. We expected High Energy Requirements cetaceans to preferentially forage in habitats of high prey biomass and/or production, where they might easily meet their high energetic needs, and Low Energy Requirements cetaceans to forage in habitats of either high or low prey biomass and/or production. Cetacean sightings were collected from dedicated aerial surveys in the South West Indian Ocean (SWIO) and French Polynesia (FP). We examined cetacean densities in relation to simulated distributions of their potential prey using Generalised Additive Models and predicted their habitats in both regions. Results supported their known diving abilities, with Delphininae mostly related to prey present in the upper layers of the water column, and Globicephalinae and sperm and beaked whales also related to prey present in deeper layers. Explained deviances ranged from 9% for sperm and beaked whales in the SWIO to 47% for Globicephalinae in FP. Delphininae and Globicephalinae appeared to select areas where high prey biomass and/or production were available at shallow depths. In contrast, sperm and beaked whales showed less clear habitat selection. Using simulated prey distributions as predictors in cetacean habitat models is crucial to understand their strategies of habitat selection in the three dimensions of the ocean.     

Plasma and urine levels of electrolytes, urea and steroid hormones involved in osmoregulation of cetaceans

Cetaceans are well adapted to their hyperosmotic environment by properly developed osmoregulatory ability. A question here is how they regulate water and mineral balances in marine habitats. In the present study, we determined blood and urine levels of various chemicals involved in osmoregulation, compared them with those in artiodactyls, and characterized the values in the whales. Blood and urine samples obtained from baleen whales of common minke (Balaenoptera acutorostrata), sei (B. borealis), and Bryde's whales (B. brydei), and toothed whales of sperm whales (Physeter macrocephalus) were analyzed for osmolality, major electrolytes, urea, steroid hormones and glucose. The urine osmolality and Na(+) concentrations in the cetaceans were much higher than those in the cattle. Furthermore, the cetaceans had 5 to 11-fold urea in plasma than the cattle, and 2 to 4-fold urea in urine. There were no significant difference in the plasma concentrations of corticosteroids between the cetaceans and the cattle. The present results indicate that the osmoregulatory parameters seem to be not affected by the reproductive stage and sex steroid hormones. The concentrations of urea in plasma and urine of the baleen whales were higher than those of the sperm whales, indicating a possibility that their osmoregulatory mechanisms may be correlated to their feeding habits. The present results suggest that cetaceans have unique osmoregulatory mechanisms by which they excrete strongly hypertonic urine to maintain fluid homeostasis in marine habitats.     

CHARACTERIZATION OF BEAKED WHALE (ZIPHIIDAE) AND SPERM WHALE (PHYSETER MACROCEPHALUS) SUMMER HABITAT IN SHELF-EDGE AND DEEPER WATERS OFF THE NORTHEAST U. S.

Sperm whales ( Physeter macrocephalus ) and beaked whales ( Mesoplodon spp. and Ziphius cavirostris ) are deep-diving cetaceans that frequent shelf-edge and Gulf Stream waters off the northeast U. S. coast. Sighting data collected during seven summer (1990, 1991, 1993, and 1995–1998) shipboard surveys were analyzed using a geographic information system to determine habitat use based on bathymetric and oceanographic features. Although sighting rates were lower for beaked whales, both taxa occupied similar habitats. Beaked whales were concentrated at the colder shelf edge, whereas sperm whales were associated with warmer off-shelf water. Mean sighting rates for both taxa were higher in canyon features, but only beaked whale sighting rates were significantly different between canyon and non-canyon habitat (Wilcoxon signed rank test P = 0.007). Within the shared habitat, the two taxa were separated at fine-scale based on oceanographic features.     

Age estimation in bowhead whales using tympanic bulla histology and baleen isotopes

Tympanic bullae and baleen plates from bowhead whales of the Western Arctic population were examined. Growth layer groups (GLGs) in the involucrum of the tympanic bone were used to estimate age of the whales, and compared to stable isotope signatures along transects of baleen plates and the involucrum. The involucrum of the tympanic bone consists of three regions that form in utero, during nursing in the first year, and during the first decades of life, respectively. Life history events, such as annual migration, are recorded in the bowhead tympanic bulla. It is likely that bone growth in the bowhead tympanic occurs during periods of high food intake, while slow or arrested growth occurs during periods of low food intake. Comparisons between numbers of GLGs in the tympanic, number of isotopic oscillations in a baleen plate, length of the baleen plate, and total whale length show correlation coefficients as high as 0.97. The tympanic GLG method is particularly useful for estimating the age of whales up to 20 yr old.     

Molecular cloning of urea transporters from the kidneys of baleen and toothed whales

Urea transport in the kidney is important for the production of concentrated urine. This process is mediated by urea transporters (UTs) encoded by two genes, UT-A (Slc14a2) and UT-B (Slc14a1). Our previous study demonstrated that cetaceans produce highly concentrated urine than terrestrial mammals, and that baleen whales showed higher concentrations of urinary urea than sperm whales. Therefore, we hypothesized that cetaceans have unique actions of UTs to maintain fluid homeostasis in marine habitat. Kidney samples of common minke (Balaenoptera acutorostrata), sei (B. borealis), Bryde's (B. brydei) and sperm whales (Physeter macrocephalus) were obtained to determine the nucleotide sequences of mRNAs encoding UT. The sequences of 2.5-kb cDNAs encode 397-amino acid proteins, which are 90-94% identical to the mammalian UT-A2s. Two putative glycosylation sites are conserved between the whales and the terrestrial mammals, whereas consensus sites for protein kinases are not completely conserved; only a single protein kinase A consensus site was identified in the whale UT-A2s. Two protein kinase C consensus sites are present in the baleen whale UT-A2s, however, a single protein kinase C consensus site was identified in the sperm whale UT-A2. These different phosphorylation sites of whale UT-A2s may result in the high concentrations of urinary urea in whales, by reflecting their urea permeability.     

Diversity versus disparity and the radiation of modern cetaceans

Modern whales are frequently described as an adaptive radiation spurred by either the evolution of various key innovations (such as baleen or echolocation) or ecological opportunity following the demise of archaic whales. Recent analyses of diversification rate shifts on molecular phylogenies raise doubts about this interpretation since they find no evidence of increased speciation rates during the early evolution of modern taxa. However, one of the central predictions of ecological adaptive radiation is rapid phenotypic diversification, and the tempo of phenotypic evolution has yet to be quantified in cetaceans. Using a time-calibrated molecular phylogeny of extant cetaceans and a morphological dataset on size, we find evidence that cetacean lineages partitioned size niches early in the evolutionary history of neocetes and that changes in cetacean size are consistent with shifts in dietary strategy. We conclude that the signature of adaptive radiations may be retained within morphological traits even after equilibrium diversity has been reached and high extinction or fluctuations in net diversification have erased any signature of an early burst of diversification in the structure of the phylogeny.     

Morphological variation among the inner ears of extinct and extant baleen whales (Cetacea: Mysticeti)

Living mysticetes (baleen whales) and odontocetes (toothed whales) differ significantly in auditory function in that toothed whales are sensitive to high‐frequency and ultrasonic sound vibrations and mysticetes to low‐frequency and infrasonic noises. Our knowledge of the evolution and phylogeny of cetaceans, and mysticetes in particular, is at a point at which we can explore morphological and physiological changes within the baleen whale inner ear. Traditional comparative anatomy and landmark‐based 3D‐geometric morphometric analyses were performed to investigate the anatomical diversity of the inner ears of extinct and extant mysticetes in comparison with other cetaceans. Principal component analyses (PCAs) show that the cochlear morphospace of odontocetes is tangential to that of mysticetes, but odontocetes are completely separated from mysticetes when semicircular canal landmarks are combined with the cochlear data. The cochlea of the archaeocete Zygorhiza kochii and early diverging extinct mysticetes plot within the morphospace of crown mysticetes, suggesting that mysticetes possess ancestral cochlear morphology and physiology. The PCA results indicate variation among mysticete species, although no major patterns are recovered to suggest separate hearing or locomotor regimes. Phylogenetic signal was detected for several clades, including crown Cetacea and crown Mysticeti, with the most clades expressing phylogenetic signal in the semicircular canal dataset. Brownian motion could not be excluded as an explanation for the signal, except for analyses combining cochlea and semicircular canal datasets for Balaenopteridae. J. Morphol. 277:1599–1615, 2016. © 2016 Wiley Periodicals, Inc.     

SITE FIDELITY, ASSOCIATIONS, AND MOVEMENTS OF CUVIER'S (ZIPHIUS CAVIROSTRIS) AND BLAINVILLE'S (MESOPLODON DENSIROSTRIS) BEAKED WHALES OFF THE ISLAND OF HAWAI'I

Although the Ziphiidae are the second-most speciose family of cetaceans, information on beaked whale species and populations has been limited by the difficulties in finding and approaching free-ranging individuals. Site fidelity, patterns of association, and movements of two species, Cuvier's ( Ziphius cavirostris ) and Blainville's ( Mesoplodon densirostris ) beaked whales, were assessed using a 21-yr photographic data set from the west coast of the island of Hawai i. Resightings of individuals of both species spanned 15 yr, suggesting long-term site fidelity to the area. Long-term resightings were documented primarily from adult females of both species. Group sizes for both species were small and most groups had only a single adult male present. For Blainville's beaked whales, repeated associations between adult females and adult males were documented for all resightings of adult males over periods from 1 to 154 d. Among adult females, although repeated associations occurred up to 9 yr apart, individuals were seen separately in intervening years. Individuals of both species seen on multiple occasions were typically documented in multiple months/seasons, suggesting they may use the study area throughout the year. Such long-term site fidelity has implications both for potential population structure and for susceptibility of beaked whale populations to anthropogenic impacts.     

The Mitochondrial Genome of the Sperm Whale and a New Molecular Reference for Estimating Eutherian Divergence Dates

Extant cetaceans are systematically divided into two suborders: Mysticeti (baleen whales) and Odontoceti (toothed whales). In this study, we have sequenced the complete mitochondrial (mt) genome of an odontocete, the sperm whale (Physeter macrocephalus), and included it in phylogenetic analyses together with the previously sequenced complete mtDNAs of two mysticetes (the fin and blue whales) and a number of other mammals, including five artiodactyls (the hippopotamus, cow, sheep, alpaca, and pig). The most strongly supported cetartiodactyl relationship was: outgroup,((pig, alpaca),((cow, sheep),(hippopotamus,(sperm whale,(baleen whales))))). As in previous analyses of complete mtDNAs, the sister-group relationship between the hippopotamus and the whales received strong support, making both Artiodactyla and Suiformes (pigs, peccaries, and hippopotamuses) paraphyletic. In addition, the analyses identified a sister-group relationship between Suina (the pig) and Tylopoda (the alpaca), although this relationship was not strongly supported. The paleontological records of both mysticetes and odontocetes extend into the Oligocene, suggesting that the mysticete and odontocete lineages diverged 32–34 million years before present (MYBP). Use of this divergence date and the complete mtDNAs of the sperm whale and the two baleen whales allowed the establishment of a new molecular reference, O/M-33, for dating other eutherian divergences. There was a general consistency between O/M-33 and the two previously established eutherian references, A/C-60 and E/R-50. Cetacean (whale) origin, i.e., the divergence between the hippopotamus and the cetaceans, was dated to ≈55 MYBP, while basal artiodactyl divergences were dated to ≥65 MYBP. Molecular estimates of Tertiary eutherian divergences were consistent with the fossil record. Received: 12 July 1999 / Accepted: 28 February 2000     

Les cétacés des gravures rupestres néolithiques de Bangu-dae (Corée du Sud) et les débuts de la chasse à la baleine dans le Pacifique nord-ouest

The Neolithic site of Bangu-dae (6000-1000 B.C.), south-east of the Korean peninsula, presents numerous rock carvings of cetaceans (n = 46, 19.9% of the figures), and especially large species. Among these two families of baleen whales (Balaenidae and Balaenopteridae) and the sperm whale can be identified. This site also presents whaling scenes suggesting that the Neolithic populations who lived along the coast of Korea were among the first to take advantage of coastal whales. All these carvings give exceptional evidence on the prehistoric beginning of whaling.     

鲸类分子系统学研究进展

综述了近年来分子生物学标记技术在鲸类系统学研究中的进展。分子生物学证据支持鲸目与有蹄类之间有较近的亲缘关系,并支持鲸类的单系起源,但鲸类不同类群(须鲸类、抹香鲸类及不包括抹香鲸类的齿鲸类)之间的系统发生关系仍存在争议。抹香鲸类到底与须鲸类还是与其它齿鲸类有更近的亲缘关系,不同的分子生物学家所得到的结果并不一致。此外,分子生物学技术还被用于解决须鲸亚目和齿鲸亚目内科间以及科内种间的系统发生关系,特别是齿鲸亚目的海豚科、鼠豚科和淡水豚类。通过分子标记技术来研究鲸类种下的遗传结构是鲸类分子系统学研究中的一个新热点,使用的标记主要是mtDNA控制区、核DNA微卫星和主要组织相容性复合体(major histo-compatibilitv complex,MHC)等。     

Ultrastructure,biomechanical and chemical properties of the vestigial dentition of a Cuvier's beaked whale

Beaked whales are poorly known cetaceans with a peculiar dentition consisting of one or two pairs of mandibular tusks, sometimes accompanied by dozens of peg-like rudimentary teeth normally concealed in the gum tissue. This study investigated the ultrastructure, chemical and biomechanical properties of vestigial teeth in a specimen of Cuvier's beaked whale deposited in the Otago Museum collections. The juvenile individual had 49 needle-like vestigial teeth in situ on the lower jaws. Scanning electron microscopy images revealed a thin layer of semi-structured prismless enamel covering the teeth. Mechanical properties values were within the range of values reported for other cetaceans, suggesting relaxed biomechanical functional demands. Chemical analyses revealed preservation of the hydroxyapatite composition despite morphological simplification and vestigial nature. The presumed high incidence of vestigial teeth in Ziphius cavirostris, the small size and the degenerate nature of these structures are consistent with their interpretation as vestiges rather than atavisms.     

The first baleen whale marine protected area proposed for Bryde's whales in the Beibu Gulf,China

The establishment of marine protected areas (MPAs) for cetaceans is an important strategy to mitigate human disturbance and protect biodiversity. Despite abundant cetacean species, there are only a few MPAs dedicated to cetacean conservation in China, all of which are for inshore dolphins. Bryde's whales, the only nearshore baleen whale population in mainland China, are conflicting with intensive human activities, yet an effective conservation strategy is lacking. This study used species distribution models to analyze distribution patterns and suitable habitats of Bryde's whales in the Beibu Gulf and proposes the first baleen whale MPA in China. Our results showed Bryde's whales have a seasonal distribution pattern in the Beibu Gulf, and that the waters around Weizhou Island and the southeastern coast of Vietnam were their core habitats. The seasonal nighttime light data indicated a negative relationship between the number of ship lights and Bryde's whale sightings and suggest that Bryde's whales might be threatened by fisheries. We proposed an MPA based on the results, suggesting that the waters within 20 km around Weizhou Island should be declared a protected area. Furthermore, we recommend that anthropogenic activities in the waters around Weizhou Island are better managed to reduce negative impacts on marine life.     

Population biology, social behavior and communication in whales and dolphins

The baleen whales differ from the toothed whales and dolphins in life history and in social organization. Even though they grow to a larger size, young baleen whales tend to develop more rapidly than dolphins and toothed whales. Except for the mother-calf bond, most groups of baleen whales are short-lived, lasting only for hours, and individual-specific associations appear to be exceptions to the norm. Most toothed whales live in more structured groups, in which young animals have a long period of dependency and social learning. The communication signals described for different cetacean species have functions suited to the interactions that predominate in their societies.     

SPERM WHALE PHYLOGENY REVISITED: ANALYSIS OF THE MORPHOLOGICAL EVIDENCE

Some recent analyses of three mitochondrial DNA regions suggest that sperm whales are the sister group to baleen whales and, therefore, the suborder Odontoceti (toothed whales) constitutes a paraphyletic group. I cladistically analyzed the available morphological data, including that from relevant fossil taxa, for all families of extant cetaceans to test this hypothesis. The results of this analysis unambiguously support a monophyletic Odontoceti including the sperm whales. All synapomorphies that support the Odontoceti node are decisive, not related to the evolution of highly correlated characters, and provide the same result regardless of what order of mammals is used as an outgroup. These numerous, anatomically diverse, and unambiguous characters make this clade one of the best-supported higher-level groupings among mammals. In addition, the fossil evidence refutes a sperm whale/baleen whale clade. Both the molecular and morphological data produce the same unrooted tree. The improper rooting of the molecular tree appears to be producing these seemingly incongruent phylogenies.     

Evolution of the Arctic Calanus complex: an Arctic marine avocado?

Before man hunted the large baleen whales to near extinction by the end of the nineteenth century, Arctic ecosystems were strongly influenced by these large predators. Their main prey were zooplankton, among which the calanoid copepod species of the genus Calanus, long considered key elements of polar marine ecosystems, are particularly abundant. These herbivorous zooplankters display a range of adaptations to the highly seasonal environments of the polar oceans, most notably extensive energy reserves and seasonal migrations to deep waters where the non-feeding season is spent in diapause. Classical work in marine ecology has suggested that slow growth, long lifespan and large body size in zooplankton are specific adaptations to life in cold waters with short and unpredictable feeding seasons. Here, we challenge this understanding and, by using an analogy from the evolutionary and contemporary history of the avocado, argue that predation pressure by the now nearly extinct baleen whales was an important driving force in the evolution of life history diversity in the Arctic Calanus complex.     

CETACEAN ABUNDANCE IN HAWAIIAN WATERS ESTIMATED FROM A SUMMER/FALL SURVEY IN 2002

Cetacean abundance is estimated for the U.S. Exclusive Economic Zone (EEZ) around the Hawaiian Islands based on a ship line‐transect survey from August to November, 2002. Sighting detection functions are estimated from this and other NOAA research surveys from 1986 to 2002 using a new, multiple‐covariate approach. Twenty‐four species were seen on this survey, including two species (Fraser's dolphin [Lagenodelphis hosei] and sei whale [Balaenoptera borealis]) that had not been previously documented to occur in Hawaiian waters. The most abundant large whales are sperm whales (Physeter macrocephalus) and Bryde's whales (Balaenoptera edeni). The most abundant delphinids are pilot whales (Globicephala macrorhynchus), rough‐toothed dolphins (Steno bredanensis), Fraser's dolphins, spotted dolphins (Stenella attenuata), and striped dolphins (Stenella coeruleoalba). Dwarf and pygmy sperm whales (Kogia sima and Kogia breviceps) and Cuvier's beaked whales (Ziphius cavirostris) are also estimated to be quite abundant. Some of the migratory baleen whales (fin whales [Balaenoptera physalus], sei whales, minke whales [B. acutorostrata], and humpback whales [Megaptera novaeangliae]) were seen only late in the survey. Abundance is estimated for 19 cetacean species. The overall density of cetaceans is low in the study area, especially for delphinids. The precision of density and abundance estimates is generally low for all species because of the low number of sightings.     

LESSONS FROM MONITORING TRENDS IN ABUNDANCE OF MARINE MAMMALS

We assessed scientists' ability to detect declines of marine mammal stocks based on recent levels of survey effort, when the actual decline is precipitous. We defined a precipitous decline as a 50% decrease in abundance in 15 yr, at which point a stock could be legally classified as "depleted" under the U.S. Marine Mammal Protection Act. We assessed stocks for three categories of cetaceans: large whales ( n = 23, most of which are listed as endangered), beaked whales ( n = 11, potentially vulnerable to anthropogenic noise), and small whales/dolphins/porpoises ( n = 69, bycatch in fisheries and important abundant predators), for two categories of pinnipeds with substantially different survey precision: counted on land ( n = 13) and surveyed on ice ( n = 5), and for a category containing polar bear and sea otter stocks ( n = 6). The percentage of precipitous declines that would not be detected as declines was 72% for large whales, 90% for beaked whales, and 78% for dolphins/porpoises, 5% for pinnipeds on land, 100% for pinnipeds on ice, and 55% for polar bears/sea otters (based on a one-tailed t -test, α= 0.05), given the frequency and precision of recent monitoring effort. We recommend alternatives to improve performance.