Observations of cetaceans in the south-east Atlantic sector of the Southern Ocean,during summer 2008

Knowledge of cetacean species composition and their distribution in the south-east Atlantic sector of the Southern Ocean is scarce. During a survey in February–March 2008, systematic whale sightings were carried out along transect lines following the 5° and 15° E meridians between 35° and 67° S. In total, 67 toothed whales and 126 baleen whales were observed. Both fin whales (four animals) and Antarctic minke whales Balaenoptera bonaerenses (three animals) in addition to 16 individuals of unidentified species were among the observed baleen whales. The dominating baleen whale species in our study was humpback whales Megaptera novaeangliae with 108 individuals observed. They occurred single or in groups up to seven individuals (N _mean = 2.5 ind) and eight of the counts were of calves. The relationship between humpback whale occurrence and environmental variables including Antarctic krill (Euphausia superba) abundance from acoustic recordings, hydrography, bathymetry and production was tested using general additive models. Only temperature increased the predictive power of the model with whale occurrence increasing with the decreasing temperature in more southern areas.     

Respiration cycle duration and seawater flux through open blowholes of humpback (Megaptera novaeangliae) and North Atlantic right (Eubalaena glacialis) whales

Little is known about the dynamics of baleen whale respiratory cycles, especially the mechanics and activity of the blowholes and their interaction with seawater. In this study, the duration of complete respiration cycles (expiration/inhalation events) were quantified for the first time in two species: North Atlantic right whale (NARW) and humpback whale (HW) using high resolution, detailed imagery from an unoccupied aerial system (UAS). The mean duration of complete respiration cycles (expiration/inhalation event) in the NARW and HW were 3.07 s (SD = 0.503, n = 15) and 2.85 s (SD = 0.581, n = 21), respectively. Furthermore, we saw no significant differences in respiration cycle duration between age and sex classes in the NARW, but significant differences were observed between age classes in the HW. The observation of seawater covering an open blowhole was also quantified, with NARW having 20% of all breaths with seawater presence versus 90% in HW. Seawater incursion has not been described previously and challenges the general consensus that water does not enter the respiratory tract in baleen whales. Prevalent seawater has implications for the analysis and interpretation of exhaled respiratory vapor/mucosa samples, as well as for the potential inhalation of oil in spills.     

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.     

Species identification and likely catch time period of whale bones from South Georgia

Skeletal remains of baleen whales killed during the onset of 20th century commercial whaling lie scattered across the shores and abandoned whaling stations of the subantarctic island of South Georgia. Here we report on genetic species identification of whale bones collected from South Georgia using standard historical DNA protocols. We amplified and sequenced short fragments of the mitochondrial DNA (mtDNA) control region from 281 available bone samples. Of these, 231 provided mtDNA sequences of sufficient quality and length (174–194 bp) for species identification: 158 bones were identified as humpback whale (Megaptera novaeangliae), 51 bones were identified as fin whale (Balaenoptera physalus), 18 bones were identified as blue whale (B. musculus), two bones were identified as sei whale (B. borealis), one bone was identified as a southern right whale (Eubalaena australis), and one bone was identified as a southern elephant seal (Mirounga leonina). The prominence of humpback, fin, and blue whale bones in the sample collection corresponds to the catch record of the early years of whaling on the island of South Georgia (pre‐1915), prior to the depletion of these populations.     

Status of the world's baleen whales

No global synthesis of the status of baleen whales has been published since the 2008 IUCN Red List assessments. Many populations remain at low numbers from historical commercial whaling, which had ceased for all but a few by 1989. Fishing gear entanglement and ship strikes are the most severe current threats. The acute and long‐term effects of anthropogenic noise and the cumulative effects of multiple stressors are of concern but poorly understood. The looming consequences of climate change and ocean acidification remain difficult to characterize. North Atlantic and North Pacific right whales are among the species listed as Endangered. Southern right, bowhead, and gray whales have been assessed as Least Concern but some subpopulations of these species ‐ western North Pacific gray whales, Chile‐Peru right whales, and Svalbard/Barents Sea and Sea of Okhotsk bowhead whales ‐ remain at low levels and are either Endangered or Critically Endangered. Eastern North Pacific blue whales have reportedly recovered, but Antarctic blue whales remain at about 1% of pre‐exploitation levels. Small isolated subspecies or subpopulations, such as northern Indian Ocean blue whales, Arabian Sea humpback whales, and Mediterranean Sea fin whales are threatened while most subpopulations of sei, Bryde's, and Omura's whales are inadequately monitored and difficult to assess.     

'Megapclicks': acoustic click trains and buzzes produced during night-time foraging of humpback whales (Megaptera novaeangliae)

Humpback whales (Megaptera novaeangliae) exhibit a variety of foraging behaviours, but neither they nor any baleen whale are known to produce broadband clicks in association with feeding, as do many odontocetes. We recorded underwater behaviour of humpback whales in a northwest Atlantic feeding area using suction-cup attached, multi-sensor, acoustic tags (DTAGs). Here we describe the first recordings of click production associated with underwater lunges from baleen whales. Recordings of over 34000 'megapclicks' from two whales indicated relatively low received levels at the tag (between 143 and 154dB re 1 microPa pp), most energy below 2kHz, and interclick intervals often decreasing towards the end of click trains to form a buzz. All clicks were recorded during night-time hours. Sharp body rolls also occurred at the end of click bouts containing buzzes, suggesting feeding events. This acoustic behaviour seems to form part of a night-time feeding tactic for humpbacks and also expands the known acoustic repertoire of baleen whales in general.     

Distribution and abundance of sei whales off the west coast of the Falkland Islands

Little information exists on the current status of Southern Hemisphere sei whales (Balaenoptera borealis). We assessed their distribution and abundance along the west coast of the Falkland Islands (southwest Atlantic) during February and March 2018, using line transect and nonsystematic surveys. Abundance estimates were generated for a single survey stratum using design- and model-based approaches. Sightings of sei whales and unidentified baleen whales (most, if not all, likely to be sei whales) occurred from the coast to the 100 m depth isobath that marked the offshore boundary of the stratum. The modeled distribution predicted highest whale densities in King George Bay and in the waters between Weddell Island and the Passage Islands. Sei whale abundance was estimated as 716 animals (CV = 0.22; 95% CI [448, 1,144]; density = 0.20 whales/km²) using the design-based approach, and 707 animals (CV = 0.11; 95% CI [566, 877]; density = 0.20 whales/km²) using the model-based approach. For sei whales and unidentified baleen whales combined, the equivalent estimates were 916 animals (CV = 0.19; 95% CI [606, 1,384]; density = 0.26 whales/km²) and 895 animals (CV = 0.074; 95% CI [777, 1,032]; density = 0.25 whales/km²). The data indicate that the Falkland Islands inner shelf region may support globally important seasonal feeding aggregations of sei whales, and potentially qualify as a Key Biodiversity Area.     

Fight or flight: antipredator strategies of baleen whales

• 1 The significance of killer whale Orcinus orca predation on baleen whales (Mysticeti) has been a topic of considerable discussion and debate in recent years. Discourse has been constrained by poor understanding of predator‐prey dynamics, including the relative vulnerability of different mysticete species and age classes to killer whales and how these prey animals avoid predation. Here we provide an overview and analysis of predatory interactions between killer whales and mysticetes, with an emphasis on patterns of antipredator responses.
• 2 Responses of baleen whales to predatory advances and attacks by killer whales appear to fall into two distinct categories, which we term the fight and flight strategies. The fight strategy consists of active physical defence, including self‐defence by single individuals, defence of calves by their mothers and coordinated defence by groups of whales. It is documented for five mysticetes: southern right whale Eubalaena australis, North Atlantic right whale Eubalaena glacialis, bowhead whale Balaena mysticetus, humpback whale Megaptera novaeangliae and grey whale Eschrichtius robustus. The flight strategy consists of rapid (20–40 km/h) directional swimming away from killer whales and, if overtaken and attacked, individuals do little to defend themselves. This strategy is documented for six species in the genus Balaenoptera.
• 3 Many aspects of the life history, behaviour and morphology of mysticetes are consistent with their antipredator strategy, and we propose that evolution of these traits has been shaped by selection for reduced predation. Fight species tend to have robust body shapes and are slow but relatively manoeuvrable swimmers. They often calve or migrate in coastal areas where proximity to shallow water provides refuge and an advantage in defence. Most fight species have either callosities (rough and hardened patches of skin) or encrustations of barnacles on their bodies, which may serve (either primarily or secondarily) as weapons or armour for defence. Flight species have streamlined body shapes for high‐speed swimming and they can sustain speeds necessary to outrun pursuing killer whales (>15–20 km/h). These species tend to favour pelagic habitats and calving grounds where prolonged escape sprints from killer whales are possible.
• 4 The rarity of observed successful attacks by killer whales on baleen whales, especially adults, may be an indication of the effectiveness of these antipredator strategies. Baleen whales likely offer low profitability to killer whales, relative to some other marine mammal prey. High‐speed pursuit of flight species has a high energetic cost and a low probability of success while attacks on fight species can involve prolonged handling times and a risk of serious injury.

     

Phenological changes in North Atlantic right whale habitat use in Massachusetts Bay

The North Atlantic right whale (Eubalaena glacialis) is one of the world's most highly endangered baleen whales, with approximately 400–450 individuals remaining. Massachusetts Bay (MB) and Cape Cod Bay (CCB) together comprise one of seven areas in the Gulf of Maine where right whales seasonally congregate. Here, we report on acoustically detected presence of right whales in MB over a nearly 6 year period, July 2007–April 2013, a time of both rapid ocean warming throughout the Gulf of Maine and apparent changes in right whale migratory dynamics. We applied an automated detection algorithm to assess hourly presence of right whale “up‐calls” in recordings from a 19‐channel acoustic array covering approximately 4,000 km² in MB. Over the survey, up‐calls were detected in 95% of 8 day periods. In each year, as expected, we observed a “peak season” of elevated up‐call detections in late winter and early spring corresponding to the season when right whales congregate to feed in CCB. However, we also saw an increase in right whale occurrence during time periods thought to be part of the “off‐season.” With the exception of 2009–2010, when acoustic presence was unusually low, the mean percent of hours in which up‐calls were detected increased every year, both during the peak season (from 38% in 2008 to 70% in 2012), and during the summer–fall season (from 2% in 2007 to 13% in 2012). Over the entire study, the peak season start date varied between 17 January and 26 February. Changes in right whale phenology in MB likely reflect broadscale changes in habitat use in other areas within the species range. This study demonstrates the value of continuous long‐term survey datasets to detect and quantify shifts in cetacean habitat use as environmental conditions change and the long‐term continued survival of right whales remains uncertain.     

Whales from space: Four mysticete species described using new VHR satellite imagery

Large‐bodied animals such as baleen whales can now be detected with very high resolution (VHR) satellite imagery, allowing for scientific studies of whales in remote and inaccessible areas where traditional survey methods are limited or impractical. Here we present the first study of baleen whales using the WorldView‐3 satellite, which has a maximum spatial resolution of 31 cm in the panchromatic band, the highest currently available to nonmilitary professionals. We manually detected, described, and counted four different mysticete species: fin whales (Balaenoptera physalus) in the Ligurian Sea, humpback whales (Megaptera novaeangliae) off Hawaii, southern right whales (Eubalaena australis) off Península Valdés, and gray whales (Eschrichtius robustus) in Laguna San Ignacio. Visual and spectral analyses were conducted for each species, their surrounding waters, and nonwhale objects (e.g., boats). We found that behavioral and morphological differences made some species more distinguishable than others. Fin and gray whales were the easiest to discern due to their contrasting body coloration with surrounding water, and their prone body position, which is proximal to the sea surface (i.e., body parallel to the sea surface). These results demonstrate the feasibility of using VHR satellite technology for monitoring the great whales.     

Tracing the spatio-temporal dynamics of endangered fin whales (<Emphasis Type="Italic">Balaenoptera physalus</Emphasis>) within baleen whale (Mysticeti) lineages: a mitogenomic perspective

To explore the spatio-temporal dynamics of endangered fin whales (Balaenoptera physalus) within the baleen whale (Mysticeti) lineages, we analyzed 148 published mitochondrial genome sequences of baleen whales. We used a Bayesian coalescent approach as well as Bayesian inferences and maximum likelihood methods. The results showed that the fin whales had a single maternal origin, and that there is a significant correlation between geographic location and evolution of global fin whales. The most recent common female ancestor of this species lived approximately 9.88 million years ago (Mya). Here, North Pacific fin whales first appeared about 7.48 Mya, followed by a subsequent divergence in Southern Hemisphere approximately 6.63 Mya and North Atlantic about 4.42 Mya. Relatively recently, approximately 1.76 and 1.42 Mya, there were two additional occurrences of North Pacific populations; one originated from the Southern Hemisphere and the other from an uncertain location. The evolutionary rate of this species was 1.002?×?10^?3 substitutions/site/My. Our Bayesian skyline plot illustrates that the fin whale population has the rapid expansion event since ~?2.5 Mya, during the Quaternary glaciation stage. Additionally, this study indicates that the fin whale has a sister group relationship with humpback whale (Meganoptera novaeangliae) within the baleen whale lineages. Of the 16 genomic regions, NADH5 showed the most powerful signal for baleen whale phylogenetics. Interestingly, fin whales have 16 species-specific amino acid residues in eight mitochondrial genes: NADH2, COX2, COX3, ATPase6, ATPase8, NADH4, NADH5, and Cytb.     

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.      

Stable isotope records of sei whale baleens from Chilean Patagonia as archives for feeding and migration behavior

Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope variations in baleen plates of sei whales (Balaenoptera borealis) stranded after a mass mortality event in Chilean Patagonia were investigated to assess potential dietary and migratory patterns. Carbon and nitrogen isotope ratios of seven baleens from six individuals were analyzed. The δ¹³C values ranged from ? 19.1 to ? 15.9‰ and the δ¹⁵N values from 8.7 to 15.4‰. Variations of up to 2.9‰ for δ¹³C and 5.3‰ for δ¹⁵N were observed within one baleen. Carbon and nitrogen isotope records of each baleen were significantly correlated and showed recurring oscillations confirmed by wavelet analyses. Oscillations slightly differed in periodicity indicating variable baleen growth rates between 10.0 and 16.5 cm/year. Food sources of the whales are discussed in terms of available isotope data for potential prey taxa and potential migratory behavior on the basis of latitudinal isotope gradients of particulate organic matter. Cyclicity could be explained by regular migrations of the sei whales from subtropical calving areas to high‐latitude foraging grounds. δ¹⁵N records of baleens differed between individuals eventually pointing to diverse feeding and migratory preferences among sei whale individuals.     

A review of cetacean occurrence in West African waters from the Gulf of Guinea to Angola

• 1 The cetacean fauna of the west coast of Africa is poorly described. Therefore, literature on the occurrence of cetacean species in the waters of 13 potential West African range states from the Gulf of Guinea to Angola was reviewed, including sighting, stranding, capture, bycatch and whaling records.
• 2 At least 28 species of cetacean were documented in the study region, comprising seven baleen whale species and 21 species of toothed whale (including at least 17 delphinid species).
• 3 Cetaceans could be broadly split into seven ecological categories, based on their distribution. A warm temperate/tropical deep‐water cetacean community dominated the study area. Cooler water from the Benguela Current influenced southern Angola (≤16°S latitude) and at least three cetacean species occurred predominantly in this region.
• 4 Only three or fewer species were confirmed in the waters of Togo, Nigeria, Cameroon and the Democratic Republic of Congo. Seventeen or more species were documented in Ghana, Gabon and Angola, where dedicated cetacean research projects have been initiated in recent years. Angola had the most diverse documented cetacean community: 28 confirmed species.
• 5 The humpback whale Megaptera novaeangliae was the most widely recorded species, and was documented in 11 (85%) countries. Sperm whales Physeter macrocephalus, Bryde's whales Balaenoptera cf. brydei, bottlenose dolphins Tursiops truncatus and Atlantic spotted dolphins Stenella frontalis were recorded in over half of the countries.

     

Adapting to a Warmer Ocean—Seasonal Shift of Baleen Whale Movements over Three Decades

Global warming poses particular challenges to migratory species, which face changes to the multiple environments occupied during migration. For many species, the timing of migration between summer and winter grounds and also within-season movements are crucial to maximise exploitation of temporarily abundant prey resources in feeding areas, themselves adapting to the warming planet. We investigated the temporal variation in the occurrence of fin (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in a North Atlantic summer feeding ground, the Gulf of St. Lawrence (Canada), from 1984 to 2010 using a long-term study of individually identifiable animals. These two sympatric species both shifted their date of arrival at a previously undocumented rate of more than 1day per year earlier over the study period thus maintaining the approximate 2-week difference in arrival of the two species and enabling the maintenance of temporal niche separation. However, the departure date of both species also shifted earlier but at different rates resulting in increasing temporal overlap over the study period indicating that this separation may be starting to erode. Our analysis revealed that the trend in arrival was strongly related to earlier ice break-up and rising sea surface temperature, likely triggering earlier primary production. The observed changes in phenology in response to ocean warming are a remarkable example of phenotypic plasticity and may partly explain how baleen whales were able to survive a number of changes in climate over the last several million years. However, it is questionable whether the observed rate of change in timing can be maintained. Substantial modification to the distribution or annual life cycle of these species might be required to keep up with the ongoing warming of the oceans.     

Diversity and duplication of DQB and DRB-like genes of the MHC in baleen whales (suborder: Mysticeti)

The molecular diversity and phylogenetic relationships of two class II genes of the baleen whale major histocompatibility complex were investigated and compared to toothed whales and out-groups. Amplification of the DQB exon 2 provided sequences showing high within-species and between-species nucleotide diversity and uninterrupted reading frames consistent with functional class II loci found in related mammals (e.g., ruminants). Cloning of amplified products indicated gene duplication in the humpback whale and triplication in the southern right whale, with average nucleotide diversity of 5.9 and 6.3%, respectively, for alleles of each species. Significantly higher nonsynonymous divergence at sites coding for peptide binding (32% for humpback and 40% for southern right) suggested that these loci were subject to positive (overdominant) selection. A population survey of humpback whales detected 23 alleles, differing by up to 21% of their inferred amino acid sequences. Amplification of the DRB exon 2 resulted in two groups of sequences. One was most similar to the DRB3 of the cow and present in all whales screened to date, including toothed whales. The second was most similar to the DRB2 of the cow and was found only in the bowhead and right whales. Both loci showed low diversity among species and apparent loss of function or altered function including interruption of reading frames. Finally, comparison of inferred protein sequence of the DRB3-like locus suggested convergence with the DQB, perhaps resulting from intergenic conversion or recombination.Electronic Supplementary Material Supplementary material is available for this article at     

Mitochondrial phylogenetics and evolution of mysticete whales

The phylogenetic relationships among baleen whales (Order: Cetacea) remain uncertain despite extensive research in cetacean molecular phylogenetics and a potential morphological sample size of over 2 million animals harvested. Questions remain regarding the number of species and the monophyly of genera, as well as higher order relationships. Here, we approach mysticete phylogeny with complete mitochondrial genome sequence analysis. We determined complete mtDNA sequences of 10 extant Mysticeti species, inferred their phylogenetic relationships, and estimated node divergence times. The mtDNA sequence analysis concurs with previous molecular studies in the ordering of the principal branches, with Balaenidae (right whales) as sister to all other mysticetes base, followed by Neobalaenidae (pygmy right whale), Eschrichtiidae (gray whale), and finally Balaenopteridae (rorquals + humpback whale). The mtDNA analysis further suggests that four lineages exist within the clade of Eschrichtiidae + Balaenopteridae, including a sister relationship between the humpback and fin whales, and a monophyletic group formed by the blue, sei, and Bryde's whales, each of which represents a newly recognized phylogenetic relationship in Mysticeti. We also estimated the divergence times of all extant mysticete species, accounting for evolutionary rate heterogeneity among lineages. When the mtDNA divergence estimates are compared with the mysticete fossil record, several lineages have molecular divergence estimates strikingly older than indicated by paleontological data. We suggest this discrepancy reflects both a large amount of ancestral polymorphism and long generation times of ancestral baleen whale populations.     

Prey preferences of sympatric fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales revealed by stable isotope mixing models

Over‐exploitation of top predators and fish stocks has altered ecosystems towards less productive systems with fewer trophic levels. In the Celtic Sea (CS), discards and bycatch levels have prompted concern about some fisheries, while fin and humpback whales are recovering from centuries of over‐exploitation. A lack of empirical evidence on the preferred diet of some predators such as whales in the CS has hindered the implementation of effective conservation measures using an ecosystem‐based approach to fisheries management. Using a Bayesian framework (SIAR), stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope mixing models were used to assign proportionate diet solutions to fin and humpback whales (skin biopsies) and putative prey items: herring (Clupea harengus), sprat (Sprattus sprattus), and krill (Meganyctiphanes norvegica and Nyctiphanes couchii) in the CS. Krill was the single most important prey item in the diet of fin whales, but one of the least important for humpback whales (albeit based on a small sample of humpback whale samples). Age 0 sprat and herring comprised a large proportion of the diet of both species, followed by older sprat (age 1–2) and older herring (age 2–4). An ecosystem based approach to fisheries management will be required in the CS if we seek effective conservation of both fin and humpback whales, and sustainable fisheries.