Morphology of nares associated with stereo-olfaction in baleen whales.

The sensory mechanisms used by baleen whales (Mysticeti) for locating ephemeral, dense prey patches in vast marine habitats are poorly understood. Baleen whales have a functional olfactory system with paired rather than single blowholes (nares), potentially enabling stereo-olfaction. Dimethyl sulfide (DMS) is an odorous gas emitted by phytoplankton in response to grazing by zooplankton. Some seabirds use DMS to locate prey, but this ability has not been demonstrated in whales. For 14 extant species of baleen whale, nares morphometrics (imagery from unoccupied aerial systems, UAS) was related to published trophic level indices using Bayesian phylogenetic mixed modelling. A significant negative relationship was found between nares width and whale trophic level (ß = -0.08, lower 95% CI = -0.13, upper 95% CI = -0.03), corresponding with a 39% increase in nares width from highest to lowest trophic level. Thus, species with nasal morphology best suited to stereo-olfaction are more zooplanktivorous. These findings provide evidence that some baleen whale species may be able to localize odorants e.g. DMS. Our results help direct future behavioural trials of olfaction in baleen whales, by highlighting the most appropriate species to study. This is a research priority, given the potential for DMS-mediated plastic ingestion by whales.

Mid-frequency song and low-frequency calls of sei whales in the Falkland Islands.

Although sei whales (Balaenoptera borealis) are distributed throughout the globe, their behaviour and vocal repertoire are poorly described. We used passive acoustic monitoring to describe the vocal behaviour of sei whales in the Falkland Islands, between December 2018 and April 2019. We isolated more than 2000 low-frequency calls for manual classification, of which 510 calls with high signal-to-noise ratio were quantitatively measured. Five categories of stereotyped call types in the 15-230 Hz range were described, some with multiple subcategories. These included some similar to previously described calls (e.g. downsweeps), but others that were novel in acoustic structure and frequency band. In the mid-frequency range, we documented a highly stereotyped, hierarchically structured and rhythmically repetitive song display. Songs were arranged in phrases with a structure composed of repetitive sub-phrases, and a diverse variety of sounds in the 1-5 kHz range. Singing commenced in late February, despite the presence of whales and calls since early December, and continued through April. These acoustic properties and behavioural characteristics indicate that this is likely a male breeding display similar to songs and singing of other balaenopterids. This is the first detailed description of a song display for sei whales, highlighting the importance of the Falkland Islands.

Western North Atlantic humpback whale fall and spring acoustic repertoire: Insight into onset and cessation of singing behavior.

Humpback whale songs have been described worldwide and studies exploring non-song vocal behavior continue to expand; however, studies on the transition periods when whales shift to and from the seasonal behavioral state of singing are lacking and may be potentially informative regarding the proximal factors controlling the onset and offset of humpback whale male singing. Acoustic recorders collected data off eastern Canada continuously from the Bay of Fundy in the fall of 2015 and near-continuously off northeast Nova Scotia in the spring of 2016. Humpback whale acoustic occurrence and behavior were identified by systematically reviewing a subset of acoustic recordings for presence before analyzing the highest quality recordings for behavior. The onset of singing in the fall was gradual over a period of about three weeks with an intermediate form, termed "song fragment," occurring prior to full songs. In comparison, singing in the spring seemed to end abruptly with few song fragments. Song fragments could be produced by juveniles learning to sing for the first time or mature males preparing for breeding activities prior to migrating to southern breeding grounds. The authors propose an alternative hypothesis that the timing and manner of transitions could be driven by physiological processes similar to those documented in songbirds.

Oceanographic barriers, divergence, and admixture: Phylogeography and taxonomy of two putative subspecies of short-finned pilot whale.

Genomic phylogeography plays an important role in describing evolutionary processes and their geographic, ecological, or cultural drivers. These drivers are often poorly understood in marine environments, which have fewer obvious barriers to mixing than terrestrial environments. Taxonomic uncertainty of some taxa (e.g., cetaceans), due to the difficulty in obtaining morphological data, can hamper our understanding of these processes. One such taxon, the short-finned pilot whale, is recognized as a single global species but includes at least two distinct morphological forms described from stranding and drive hunting in Japan, the "Naisa" and "Shiho" forms. Using samples (n = 735) collected throughout their global range, we examine phylogeographic patterns of divergence by comparing mitogenomes and nuclear SNP loci. Our results suggest three types within the species: an Atlantic Ocean type, a western/central Pacific and Indian Ocean (Naisa) type, and an eastern Pacific Ocean and northern Japan (Shiho) type. mtDNA control region differentiation indicates these three types form two subspecies, separated by the East Pacific Barrier: Shiho short-finned pilot whale, in the eastern Pacific Ocean and northern Japan, and Naisa short-finned pilot whale, throughout the remainder of the species' distribution. Our data further indicate two diverging populations within the Naisa subspecies, in the Atlantic Ocean and western/central Pacific and Indian Oceans, separated by the Benguela Barrier off South Africa. This study reveals a process of divergence and speciation within a globally-distributed, mobile marine predator, and indicates the importance of the East Pacific Barrier to this evolutionary process.

Songbird dynamics under the sea: acoustic interactions between humpback whales suggest song mediates male interactions.

The function of song has been well studied in numerous taxa and plays a role in mediating both intersexual and intrasexual interactions. Humpback whales are among few mammals who sing, but the role of sexual selection on song in this species is poorly understood. While one predominant hypothesis is that song mediates male-male interactions, the mechanism by which this may occur has never been explored. We applied metrics typically used to assess songbird interactions to examine song sequences and movement patterns of humpback whale singers. We found that males altered their song presentation in the presence of other singers; focal males increased the rate at which they switched between phrase types (p = 0.005), and tended to increase the overall evenness of their song presentation (p = 0.06) after a second male began singing. Two-singer dyads overlapped their song sequences significantly more than expected by chance. Spatial analyses revealed that change in distance between singers was related to whether both males kept singing (p = 0.012), with close approaches leading to song cessation. Overall, acoustic interactions resemble known mechanisms of mediating intrasexual interactions in songbirds. Future work should focus on more precisely resolving how changes in song presentation may be used in competition between singing males.

Continuous movement behavior of humpback whales during the breeding season in the southwest Indian Ocean: on the road again!

BACKGROUND: Humpback whales are known to undertake long-distance migration between feeding and breeding sites, but their movement behavior within their breeding range is still poorly known. Satellite telemetry was used to investigate movement of humpback whales during the breeding season and provide further understanding of the breeding ecology and sub-population connectivity within the southwest Indian Ocean (SWIO). Implantable Argos satellite tags were deployed on 15 whales (7 males and 6 females) during the peak of the breeding season in Reunion Island. A switching-state-space model was applied to the telemetry data, in order to discriminate between "transiting" and "localized" movements, the latter of which relates to meandering behavior within putative breeding habitats, and a kernel density analysis was used to assess the spatial scale of the main putative breeding sites. RESULTS: Whales were tracked for up to 71 days from 31/07/2013 to 16/10/2013. The mean transmission duration was 25.7 days and the mean distance travelled was 2125.8 km. The tracks showed consistent movement of whales from Reunion to Madagascar, demonstrating a high level of connectivity between the two sub-regions, and the use of yet unknown breeding sites such as underwater seamounts (La Perouse) and banks (Mascarene Plateau). A localized movement pattern occurred in distinct bouts along the tracks, suggesting that whales were involved in breeding activity for 4.3 consecutive days on average, after which they resume transiting for an average of 6.6 days. Males visited several breeding sites within the SWIO, suggesting for the first time a movement strategy at a basin scale to maximize mating. Unexpectedly, females with calf also showed extensive transiting movement, while they engaged in localized behavior mainly off Reunion and Sainte-Marie (East Madagascar). CONCLUSIONS: The results indicated that whales from Reunion do not represent a discrete population. Discrete breeding sites were identified, thereby highlighting priority areas for conservation. The study is a first attempt to quantify movement of humpback whales within the southwestern Indian Ocean breeding range. We demonstrate a wandering behavior with stopovers at areas that likely represent key breeding habitat, a strategy which may enhance likelihood of individual reproductive success.

Multiple processes drive genetic structure of humpback whale (Megaptera novaeangliae) populations across spatial scales.

Elucidating patterns of population structure for species with complex life histories, and disentangling the processes driving such patterns, remains a significant analytical challenge. Humpback whale (Megaptera novaeangliae) populations display complex genetic structures that have not been fully resolved at all spatial scales. We generated a data set of nuclear markers for 3575 samples spanning the seven breeding stocks and substocks found in the South Atlantic and western and northern Indian Oceans. For the total sample, and males and females separately, we assessed genetic diversity, tested for genetic differentiation between putative populations and isolation by distance, estimated the number of genetic clusters without a priori population information and estimated rates of gene flow using maximum-likelihood and Bayesian approaches. At the ocean basin scale, structure is governed by geographical distance (IBD P < 0.05) and female fidelity to breeding areas, in line with current understanding of the drivers of broadscale population structure. Consistent with previous studies, the Arabian Sea breeding stock was highly genetically differentiated (FST 0.034-0.161; P < 0.01 for all comparisons). However, the breeding stock boundary between west South Africa and east Africa was more porous than expected based on genetic differentiation, cluster and geneflow analyses. Instances of male fidelity to breeding areas and relatively high rates of dispersal for females were also observed between the three substocks in the western Indian Ocean. The relationships between demographic units and current management boundaries may have ramifications for assessments of the status and continued protections of populations still in recovery from commercial whaling.

Influence of environmental parameters on movements and habitat utilization of humpback whales (Megaptera novaeangliae) in the Madagascar breeding ground.

Assessing the movement patterns and key habitat features of breeding humpback whales is a prerequisite for the conservation management of this philopatric species. To investigate the interactions between humpback whale movements and environmental conditions off Madagascar, we deployed 25 satellite tags in the northeast and southwest coast of Madagascar. For each recorded position, we collated estimates of environmental variables and computed two behavioural metrics: behavioural state of 'transiting' (consistent/directional) versus 'localized' (variable/non-directional), and active swimming speed (i.e. speed relative to the current). On coastal habitats (i.e. bathymetry < 200 m and in adjacent areas), females showed localized behaviour in deep waters (191 ± 20 m) and at large distances (14 ± 0.6 km) from shore, suggesting that their breeding habitat extends beyond the shallowest waters available close to the coastline. Males' active swimming speed decreased in shallow waters, but environmental parameters did not influence their likelihood to exhibit localized movements, which was probably dominated by social factors instead. In oceanic habitats, both males and females showed localized behaviours in shallow waters and favoured high chlorophyll-a concentrations. Active swimming speed accounts for a large proportion of observed movement speed; however, breeding humpback whales probably exploit prevailing ocean currents to maximize displacement. This study provides evidence that coastal areas, generally subject to strong human pressure, remain the core habitat of humpback whales off Madagascar. Our results expand the knowledge of humpback whale habitat use in oceanic habitat and response to variability of environmental factors such as oceanic current and chlorophyll level.

Omura's whales (Balaenoptera omurai) off northwest Madagascar: ecology, behaviour and conservation needs.

The Omura's whale (Balaenoptera omurai) was described as a new species in 2003 and then soon after as an ancient lineage basal to a Bryde's/sei whale clade. Currently known only from whaling and stranding specimens primarily from the western Pacific and eastern Indian Oceans, there exist no confirmed field observations or ecological/behavioural data. Here we present, to our knowledge, the first genetically confirmed documentation of living Omura's whales including descriptions of basic ecology and behaviour from northwestern Madagascar. Species identification was confirmed through molecular phylogenetic analyses of biopsies collected from 18 adult animals. All individuals shared a single haplotype in a 402 bp sequence of mtDNA control region, suggesting low diversity and a potentially small population. Sightings of 44 groups indicated preference for shallow-water shelf habitat with sea surface temperature between 27.4°C and 30.2°C. Frequent observations were made of lunge feeding, possibly on zooplankton. Observations of four mothers with young calves, and recordings of a song-like vocalization probably indicate reproductive behaviour. Social organization consisted of loose aggregations of predominantly unassociated single individuals spatially and temporally clustered. Photographic recapture of a female re-sighted the following year with a young calf suggests site fidelity or a resident population. Our results demonstrate that the species is a tropical whale without segregation of feeding and breeding habitat, and is probably non-migratory; our data extend the range of this poorly studied whale into the western Indian Ocean. Exclusive range restriction to tropical waters is rare among baleen whale species, except for the various forms of Bryde's whales and Omura's whales. Thus, the discovery of a tractable population of Omura's whales in the tropics presents an opportunity for understanding the ecological factors driving potential convergence of life-history patterns with the distantly related Bryde's whales.

Assessment of the Conservation Status of the Indian Ocean Humpback Dolphin (Sousa plumbea) Using the IUCN Red List Criteria.

Indian Ocean humpback dolphins (Sousa plumbea) are obligate shallow-water dolphins that occur exclusively in the near-shore waters of the Indian Ocean, from South Africa to the Bay of Bengal. They have a narrow habitat preference, restricted distribution and do not appear very abundant across any part of their range. There is no estimate of total species abundance; all populations that have been quantitatively evaluated have been small in size, usually fewer than 200 individuals. Fishing, dredging, land reclamation, construction blasting, port and harbour construction, pollution, boat traffic and other coastal development activities all occur, or are concentrated within, humpback dolphin habitat and threaten their survival. Although data are far from sufficient to make a rigorous quantitative assessment of population trends for this species, the scale of threats is large enough over a significant enough portion of the range to suspect or infer a decline of at least 50% over three generations, which qualifies it for listing on the IUCN Red List as Endangered. The issue primarily responsible is incidental mortality in fisheries, but the loss and degradation of habitat is likely a contributing factor. None of the threats have been adequately addressed in any part of the species' range, even though threat levels are increasing virtually everywhere.

Ecology and Conservation Status of Indian Ocean Humpback Dolphins (Sousa plumbea) in Madagascar.

The Indian Ocean humpback dolphin (Sousa plumbea) has been studied in several range states in the Southwest Indian Ocean, however little information exists on populations in Madagascar. Here, we review available literature and describe a study on S. plumbea conducted between 2004 and 2013 on the west coast of Madagascar, involving boat-based field surveys in the southwest and northwest regions, and interview surveys with local fishers from villages along most of the west coast. Field surveys in the southwest region of Anakao/St. Augustine Bay revealed low encounter rates and mean group size, and markedly declining trends in both from 1999 to 2013. Conversely, in the northwest region around Nosy Be and Nosy Iranja, encounter rates were higher, as were mean group sizes, suggesting an apparently more abundant and less impacted population. Interview surveys revealed by-catch of coastal dolphins along the entire west coast, including S. plumbea, as well as other species. Directed hunting, including drive hunts of groups of dolphins, was reported primarily in the southern regions, in the range of the Vezo Malagasy ethnicity; however, there was evidence of hunting starting in one area in the northwest, where hunting dolphins is normally considered taboo for the predominant Sakalava ethnicity. Thus, the conservation status of S. plumbea in Madagascar appears to be spatially heterogeneous, with some areas where the local population is apparently more impacted than others. Conservation measures are recommended to mitigate further decline in the southwest of Madagascar, while protecting habitat and ensuring resilience in the northwest.

Seismic surveys negatively affect humpback whale singing activity off northern Angola.

Passive acoustic monitoring was used to document the presence of singing humpback whales off the coast of Northern Angola, and opportunistically test for the effect of seismic survey activity in the vicinity on the number of singing whales. Two Marine Autonomous Recording Units (MARUs) were deployed between March and December 2008 in the offshore environment. Song was first heard in mid June and continued through the remaining duration of the study. Seismic survey activity was heard regularly during two separate periods, consistently throughout July and intermittently in mid-October/November. Numbers of singers were counted during the first ten minutes of every hour for the period from 24 May to 1 December, and Generalized Additive Mixed Models (GAMMs) were used to assess the effect of survey day (seasonality), hour (diel variation), moon phase and received levels of seismic survey pulses (measured from a single pulse during each ten-minute sampled period) on singer number. Application of GAMMs indicated significant seasonal variation, which was the most pronounced effect when assessing the full dataset across the entire season (p<0.001); however seasonality almost entirely dropped out of top-ranked models when applied to a reduced dataset during the July period of seismic survey activity. Diel variation was significant in both the full and reduced datasets (from p<0.01 to p<0.05) and often included in the top-ranked models. The number of singers significantly decreased with increasing received level of seismic survey pulses (from p<0.01 to p<0.05); this explanatory variable was included among the top ranked models for one MARU in the full dataset and both MARUs in the reduced dataset. This suggests that the breeding display of humpback whales is disrupted by seismic survey activity, and thus merits further attention and study, and potentially conservation action in the case of sensitive breeding populations.

Ancient mitochondrial M haplogroups identified in the Southwest Pacific.

Based on whole mtDNA sequencing of 14 samples from Northern Island Melanesia, we characterize three formerly unresolved branches of macrohaplogroup M that we call haplogroups M27, M28, and M29. Our 1,399 mtDNA control region sequences and a literature search indicate these haplogroups have extremely limited geographical distributions. Their coding region variation suggests diversification times older than the estimated date for the initial settlement of Northern Island Melanesia. This finding indicates that they were among the earliest mtDNA variants to appear in these islands or in the ancient continent of Sahul. These haplogroups from Northern Island Melanesia extend the existing schema for macrohaplogroup M, with many independent branches distributed across Asia, East Africa, Australia, and Near Oceania.