Interchange of Southern Hemisphere humpback whales across the South Atlantic Ocean.

The cosmopolitan distribution of humpback whales (Megaptera novaeangliae) is largely driven by migrations between winter low-latitude breeding grounds and summer high-latitude feeding grounds. Southern Hemisphere humpback whales faced intensive exploitation during the whaling eras and recently show evidence of population recovery. Gene flow and shared song indicate overlap between the western (A) and eastern (B1, B2) Breeding Stocks in the South Atlantic and Indian Oceans (C1). Here, we investigated photo-identification evidence of population interchange using images of individuals photographed during boat-based tourism and research in Brazil and South Africa from 1989 to 2022. Fluke images were uploaded to Happywhale, a global digital database for marine mammal identification. Six whales were recaptured between countries from 2002 to 2021 with resighting intervals ranging from 0.76 to 12.92 years. Four whales originally photographed off Abrolhos Bank, Brazil were photographed off the Western Cape, South Africa (feeding grounds for B2). Two whales originally photographed off the Western Cape were photographed off Brazil, one traveling to the Eastern Cape in the Southwestern Indian Ocean (a migration corridor for C1) before migrating westward to Brazil. These findings photographically confirm interchange of humpback whales across the South Atlantic and Indian Oceans and the importance of international collaboration to understand population boundaries.

Oceanographic anomalies coinciding with humpback whale super-group occurrences in the Southern Benguela.

Seasonal feeding behaviour of humpback whales (Megaptera novaeangliae) has been observed in the coastal waters of the Southern Benguela where the species has been observed forming super-groups during the austral spring in recent years since 2011. Super-groups are unprecedented densely-packed aggregations of between 20 and 200 individuals in low-latitude waters and their occurrences indicate possible changes in feeding behaviour of the species. We accessed published data on super-groups occurrence in the study area in 2011, 2014 and 2015, and investigated oceanographic drivers that support prey availability in this region. We found that enhanced primary production is a necessary but not sufficient condition for super-groups to occur. Positive chlorophyll anomalies occurring one month prior to the super-group occurrences were identified, but only a concurrent significantly reduced water volume export from the region throughout October were conducive to the aggregations in the specific years. Hydrodynamic model results attributed the anomalous decreased volume export to the strength and orientation of the Goodhope Jet and associated eddy activity. The combination of random enhanced primary production typical of the region and emerging anomalous conditions of reduced water export in October since 2011 resulted in favourable food availability leading to the unique humpback whale aggregations. The novelty of this grouping behaviour is indicative of the lack of such oceanographic conditions in the past. Given the recency of the events, it is difficult to attribute this reduction in ocean transport to climatic regime shifts, and the origin should be likely investigated in the distant water mass interaction with the greater Agulhas system rather than in local intensifications of the upwelling conditions. A positive trend in the humpback whale population abundance points to the need to monitor the exposure of the species to the changing climate conditions.

Species distribution modelling of Bryde's whales, humpback whales, southern right whales, and sperm whales in the southern African region to inform their conservation in expanding economies.

In southern African waters, information about species distribution and habitat preferences of many cetacean species is limited, despite the recent economic growth that may affect them. We determined the relative importance of eight environmental variables (bathymetry, distance to shore, slope, chlorophyll-a, salinity, eastwards sea water velocity, northwards sea water velocity and sea surface temperature) as drivers of seasonal habitat preferences of Bryde's whales (Balaenoptera brydei), humpback whales (Megaptera novaeangliae), southern right whales (Eubalaena australis) and sperm whales (Physeter macrocephalus). Using presence only data from multiple sources, we constructed predictive species distribution models (SDMs) consisting of ensembles of seven algorithms for these species during both summer and winter. Predicted distribution for all cetaceans was high in southern Africa and, in particular, within the South African Exclusive Economic Zone (EEZ). Predictive models indicated a more pronounced seasonal variation for humpback, sperm and southern right whales than for Bryde's whales. Southern right whales occurred closer to shore during winter, humpback whales were more likely to occur along the east coast in winter and the west coast in summer, and sperm whales were more concentrated off the shelf in winter. Our study shows that ensemble models using historical, incidental and scientific data, in conjunction with modern environmental variables, can provide baseline knowledge on important environmental drivers of cetacean distribution for conservation purposes. Results of this study can further be used to help develop marine spatial plans and identify important marine mammal areas.

Inter-annual decrease in pulse rate and peak frequency of Southeast Pacific blue whale song types.

A decrease in the frequency of two southeast Pacific blue whale song types was examined over decades, using acoustic data from several different sources in the eastern Pacific Ocean ranging between the Equator and Chilean Patagonia. The pulse rate of the song units as well as their peak frequency were measured using two different methods (summed auto-correlation and Fourier transform). The sources of error associated with each measurement were assessed. There was a linear decline in both parameters for the more common song type (southeast Pacific song type n.2) between 1997 to 2017. An abbreviated analysis, also showed a frequency decline in the scarcer southeast Pacific song type n.1 between 1970 to 2014, revealing that both song types are declining at similar rates. We discussed the use of measuring both pulse rate and peak frequency to examine the frequency decline. Finally, a comparison of the rates of frequency decline with other song types reported in the literature and a discussion on the reasons of the frequency shift are presented.

The recovery of North Atlantic right whales, Eubalaena glacialis, has been constrained by human-caused mortality.

North Atlantic right whales (NARW), Eubalaena glacialis, were nearly exterminated by historical whaling. Their abundance slowly increased up until 2010, to a maximum of fewer than 500 whales, and since then they have been in decline. We assessed the extent to which the relatively slow increase demonstrated by NARW was intrinsic, and how much could be due to anthropogenic impacts. In order to do so, we first compared calf counts of three populations of Southern right whales (SRW), E. australis, with that of NARW, over the period 1992-2016. By this index, the annual rate of increase of NARW was approximately one-third of that of SRW. Next we constructed a population projection model for female NARW, using the highest annual survival estimates available from recent mark-resight analysis, and assuming a four-year calving interval. The model results indicated an intrinsic rate of increase of 4% per year, approximately twice that observed, and that adult female mortality is the main factor influencing this rate. Necropsy records demonstrate that anthropogenic mortality is the primary cause of known mortality of NARW. Anthropogenic mortality and morbidity has limited the recovery of NARW, and baseline conditions prior to their recent decline were already jeopardizing NARW recovery.

A hydrodynamically active flipper-stroke in humpback whales.

A central paradigm of aquatic locomotion is that cetaceans use fluke strokes to power their swimming while relying on lift and torque generated by the flippers to perform maneuvers such as rolls, pitch changes and turns [1]. Compared to other cetaceans, humpback whales (Megaptera novaeangliae) have disproportionately large flippers with added structural features to aid in hydrodynamic performance [2,3]. Humpbacks use acrobatic lunging maneuvers to attack dense aggregations of krill or small fish, and their large flippers are thought to increase their maneuverability and thus their ability to capture prey. Immediately before opening their mouths, humpbacks will often rapidly move their flippers, and it has been hypothesized that this movement is used to corral prey [4,5] or to generate an upward pitching moment to counteract the torque caused by rapid water engulfment [6]. Here, we demonstrate an additional function for the rapid flipper movement during lunge feeding: the flippers are flapped using a complex, hydrodynamically active stroke to generate lift and increase propulsive thrust. We estimate that humpback flipper-strokes are capable of producing large forward oriented forces, which may be used to enhance lunge feeding performance. This behavior is the first observation of a lift-generating flipper-stroke for propulsion cetaceans and provides an additional function for the uniquely shaped humpback whale flipper.

Humpback whale "super-groups" - A novel low-latitude feeding behaviour of Southern Hemisphere humpback whales (Megaptera novaeangliae) in the Benguela Upwelling System.

Southern Hemisphere humpback whales (Megaptera novaeangliae) generally undertake annual migrations from polar summer feeding grounds to winter calving and nursery grounds in subtropical and tropical coastal waters. Evidence for such migrations arises from seasonality of historic whaling catches by latitude, Discovery and natural mark returns, and results of satellite tagging studies. Feeding is generally believed to be limited to the southern polar region, where Antarctic krill (Euphausia superba) has been identified as the primary prey item. Non-migrations and / or suspended migrations to the polar feeding grounds have previously been reported from a summer presence of whales in the Benguela System, where feeding on euphausiids (E. lucens), hyperiid amphipods (Themisto gaudichaudii), mantis shrimp (Pterygosquilla armata capensis) and clupeid fish has been described. Three recent research cruises (in October/November 2011, October/November 2014 and October/November 2015) identified large tightly-spaced groups (20 to 200 individuals) of feeding humpback whales aggregated over at least a one-month period across a 220 nautical mile region of the southern Benguela System. Feeding behaviour was identified by lunges, strong milling and repetitive and consecutive diving behaviours, associated bird and seal feeding, defecations and the pungent "fishy" smell of whale blows. Although no dedicated prey sampling could be carried out within the tightly spaced feeding aggregations, observations of E. lucens in the region of groups and the full stomach contents of mantis shrimp from both a co-occurring predatory fish species (Thyrsites atun) and one entangled humpback whale mortality suggest these may be the primary prey items of at least some of the feeding aggregations. Reasons for this recent novel behaviour pattern remain speculative, but may relate to increasing summer humpback whale abundance in the region. These novel, predictable, inter-annual, low latitude feeding events provide considerable potential for further investigation of Southern Hemisphere humpback feeding behaviours in these relatively accessible low-latitude waters.

Modelling the effects of environmental conditions on the acoustic occurrence and behaviour of Antarctic blue whales.

Harvested to perilously low numbers by commercial whaling during the past century, the large scale response of Antarctic blue whales Balaenoptera musculus intermedia to environmental variability is poorly understood. This study uses acoustic data collected from 586 sonobuoys deployed in the austral summers of 1997 through 2009, south of 38°S, coupled with visual observations of blue whales during the IWC SOWER line-transect surveys. The characteristic Z-call and D-call of Antarctic blue whales were detected using an automated detection template and visual verification method. Using a random forest model, we showed the environmental preferences pattern, spatial occurrence and acoustic behaviour of Antarctic blue whales. Distance to the southern boundary of the Antarctic Circumpolar Current (SBACC), latitude and distance from the nearest Antarctic shores were the main geographic predictors of blue whale call occurrence. Satellite-derived sea surface height, sea surface temperature, and productivity (chlorophyll-a) were the most important environmental predictors of blue whale call occurrence. Call rates of D-calls were strongly predicted by the location of the SBACC, latitude and visually detected number of whales in an area while call rates of Z-call were predicted by the SBACC, latitude and longitude. Satellite-derived sea surface height, wind stress, wind direction, water depth, sea surface temperatures, chlorophyll-a and wind speed were important environmental predictors of blue whale call rates in the Southern Ocean. Blue whale call occurrence and call rates varied significantly in response to inter-annual and long term variability of those environmental predictors. Our results identify the response of Antarctic blue whales to inter-annual variability in environmental conditions and highlighted potential suitable habitats for this population. Such emerging knowledge about the acoustic behaviour, environmental and habitat preferences of Antarctic blue whales is important in improving the management and conservation of this highly depleted species.

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.

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.

The world's most isolated and distinct whale population? Humpback whales of the Arabian Sea.

A clear understanding of population structure is essential for assessing conservation status and implementing management strategies. A small, non-migratory population of humpback whales in the Arabian Sea is classified as "Endangered" on the IUCN Red List of Threatened Species, an assessment constrained by a lack of data, including limited understanding of its relationship to other populations. We analysed 11 microsatellite markers and mitochondrial DNA sequences extracted from 67 Arabian Sea humpback whale tissue samples and compared them to equivalent datasets from the Southern Hemisphere and North Pacific. Results show that the Arabian Sea population is highly distinct; estimates of gene flow and divergence times suggest a Southern Indian Ocean origin but indicate that it has been isolated for approximately 70,000 years, remarkable for a species that is typically highly migratory. Genetic diversity values are significantly lower than those obtained for Southern Hemisphere populations and signatures of ancient and recent genetic bottlenecks were identified. Our findings suggest this is the world's most isolated humpback whale population, which, when combined with low population abundance estimates and anthropogenic threats, raises concern for its survival. We recommend an amendment of the status of the population to "Critically Endangered" on the IUCN Red List.

Chilean blue whales as a case study to illustrate methods to estimate abundance and evaluate conservation status of rare species.

Often abundance of rare species cannot be estimated with conventional design-based methods, so we illustrate with a population of blue whales (Balaenoptera musculus) a spatial model-based method to estimate abundance. We analyzed data from line-transect surveys of blue whales off the coast of Chile, where the population was hunted to low levels. Field protocols allowed deviation from planned track lines to collect identification photographs and tissue samples for genetic analyses, which resulted in an ad hoc sampling design with increased effort in areas of higher densities. Thus, we used spatial modeling methods to estimate abundance. Spatial models are increasingly being used to analyze data from surveys of marine, aquatic, and terrestrial species, but estimation of uncertainty from such models is often problematic. We developed a new, broadly applicable variance estimator that showed there were likely 303 whales (95% CI 176-625) in the study area. The survey did not span the whales' entire range, so this is a minimum estimate. We estimated current minimum abundance relative to pre-exploitation abundance (i.e., status) with a population dynamics model that incorporated our minimum abundance estimate, likely population growth rates from a meta-analysis of rates of increase in large baleen whales, and two alternative assumptions about historic catches. From this model, we estimated that the population was at a minimum of 16.5% (95% CI 7.3-34.4%) of pre-exploitation levels in 1998 under one catch assumption and 12.4% (CI 5.4-26.3%) of pre-exploitation levels under the other. Thus, although Chilean blue whales are probably still at a small fraction of pre-exploitation abundance, even these minimum abundance estimates demonstrate that their status is better than that of Antarctic blue whales, which are still <1% of pre-exploitation population size. We anticipate our methods will be broadly applicable in aquatic and terrestrial surveys for rarely encountered species, especially when the surveys are intended to maximize encounter rates and estimate abundance.

Population structure of humpback whales from their breeding grounds in the South Atlantic and Indian Oceans.

Although humpback whales are among the best-studied of the large whales, population boundaries in the Southern Hemisphere (SH) have remained largely untested. We assess population structure of SH humpback whales using 1,527 samples collected from whales at fourteen sampling sites within the Southwestern and Southeastern Atlantic, the Southwestern Indian Ocean, and Northern Indian Ocean (Breeding Stocks A, B, C and X, respectively). Evaluation of mtDNA population structure and migration rates was carried out under different statistical frameworks. Using all genetic evidence, the results suggest significant degrees of population structure between all ocean basins, with the Southwestern and Northern Indian Ocean most differentiated from each other. Effective migration rates were highest between the Southeastern Atlantic and the Southwestern Indian Ocean, followed by rates within the Southeastern Atlantic, and the lowest between the Southwestern and Northern Indian Ocean. At finer scales, very low gene flow was detected between the two neighbouring sub-regions in the Southeastern Atlantic, compared to high gene flow for whales within the Southwestern Indian Ocean. Our genetic results support the current management designations proposed by the International Whaling Commission of Breeding Stocks A, B, C, and X as four strongly structured populations. The population structure patterns found in this study are likely to have been influenced by a combination of long-term maternally directed fidelity of migratory destinations, along with other ecological and oceanographic features in the region.

Discovery of a blue whale feeding and nursing ground in southern Chile.

After the extensive exploitation that reduced the Southern Hemisphere blue whale (Balaenoptera musculus) populations to less than 3% of its original numbers, studies on its recovery have been compounded by the inaccessibility of most populations and the extensive migrations between low and high latitudes, thus ensuring that knowledge about blue whale ecology and status remains limited. We report the recent discovery of, arguably, the most important blue whale feeding and nursing ground known to date in the Southern Hemisphere, which is located near the fjords off southern Chile. Through aerial and marine surveys (n = 7) 47 groups, comprising 153 blue whales including at least 11 mother-calf pairs, were sighted during the austral summer and early autumn of 2003. The implications of this discovery on the biological understanding and conservation of this endangered species are discussed.