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.

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.

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.