Investigating the Relationship Between Body Shape and Life History Traits in Toothed Whales: Can Body Shape Predict Fast-Slow Life Histories?

A widespread pattern in vertebrate life-history evolution is for species to evolve towards either fast or slow life histories; however, the underlying causes of this pattern remain unclear. Toothed whales (Odontoceti) are a diverse group with a range of body sizes and life histories, making them an ideal model to investigate potential drivers of this dichotomy. Using ancestral reconstruction, we identified that certain groups of odontocetes evolved more-streamlined, presumably faster, body shapes around the same time that killer whales (Orcinus orca) evolved into whale predators approximately 1 Mya during the Pleistocene. This suggests that the evolution of a streamlined body shape may have been an adaptation to escape killer whale predation, leading to longer life-history events. To test this hypothesis, we performed a cluster analysis of odontocete whales and confirmed the dual pattern of life-history traits, with one group referred to as 'reproducers' characterized by early age of maturity, short gestation, short interbirth interval, and short lifespan, and the other group referred to as 'bet-hedgers' exhibiting the opposite pattern. However, we found that life history grouping was relatively unrelated to whale shape (i.e., more streamlined or less streamlined). Therefore, we incorporated principal component results into mixed effects models, and the model results indicated that body shape was positively related to neonate length (a measure of investment in progeny), but not significantly related to the temporal life-history traits. Thus, whale body shape is not a sufficient explanation for the evolution of fast-slow life histories in odontocete whales. Supplementary Information: The online version contains supplementary material available at 10.1007/s11692-023-09605-4.

Correction: Evidence of molting and the function of "rock-nosing" behavior in bowhead whales in the eastern Canadian Arctic.

[This corrects the article DOI: 10.1371/journal.pone.0186156.].

Evidence of molting and the function of "rock-nosing" behavior in bowhead whales in the eastern Canadian Arctic.

Bowhead whales (Balaena mysticetus) have a nearly circumpolar distribution, and occasionally occupy warmer shallow coastal areas during summertime that may facilitate molting. However, relatively little is known about the occurrence of molting and associated behaviors in bowhead whales. We opportunistically observed whales in Cumberland Sound, Nunavut, Canada with skin irregularities consistent with molting during August 2014, and collected a skin sample from a biopsied whale that revealed loose epidermis and sloughing. During August 2016, we flew a small unmanned aerial system (sUAS) over whales to take video and still images to: 1) determine unique individuals; 2) estimate the proportion of the body of unique individuals that exhibited sloughing skin; 3) determine the presence or absence of superficial lines representative of rock-rubbing behavior; and 4) measure body lengths to infer age-class. The still images revealed that all individuals (n = 81 whales) were sloughing skin, and that nearly 40% of them had mottled skin over more than two-thirds of their bodies. The video images captured bowhead whales rubbing on large rocks in shallow, coastal areas-likely to facilitate molting. Molting and rock rubbing appears to be pervasive during late summer for whales in the eastern Canadian Arctic.

Beluga whale summer habitat associations in the Nelson River estuary, western Hudson Bay, Canada.

To understand beluga whale (Delphinapterus leucas) estuarine use in the Nelson River estuary, southwest Hudson Bay, we recorded and examined beluga movements and habitat associations for the July through August period in 2002-2005. We compared locations of belugas fitted with satellite transmitters ("tags") (2002-2005) and aerial-surveyed (2003 and 2005) belugas for years of differing freshwater flow from the Nelson River which is influenced by hydroelectric activity. Using the beluga telemetry location data, we estimated an early August behavioral shift in beluga distribution patterns from local estuarine use to a progressively more migratory behavior away from the estuary. The timing of this shift in behavior was also apparent in results of beluga aerial surveys from the 1940s-1960s, despite environmental changes including later freeze-up and warming ocean temperatures. Overall, during the higher than average discharge ("wet") year of 2005, the three tagged belugas ranged farther from the Nelson River but not farther from the nearest shore along southwestern Hudson Bay, compared to the 10 tagged belugas tracked during the "dry" years of 2002-2004 with below average discharges. Aerial survey data for 2003 and 2005 display a similar dry vs. wet year shift in spatial patterns, with no significant change in overall density of belugas within the study area. In the Nelson estuary, proximity to the fresh-salt water mixing area may be more important than the shallow waters of the upper estuary. Killer whales (Orcinus orca) were observed in the Churchill area (200 km northwest) during each year of study, 2002-05, and belugas may benefit from the proximity to shallow estuary waters that provide protection from the larger-bodied predator. Study results contribute to an understanding of the influence of environmental variation on how and why belugas use estuaries although considerable uncertainties exist and additional research is required.

Sustained disruption of narwhal habitat use and behavior in the presence of Arctic killer whales.

Although predators influence behavior of prey, analyses of electronic tracking data in marine environments rarely consider how predators affect the behavior of tracked animals. We collected an unprecedented dataset by synchronously tracking predator (killer whales, [Formula: see text] = 1; representing a family group) and prey (narwhal, [Formula: see text] = 7) via satellite telemetry in Admiralty Inlet, a large fjord in the Eastern Canadian Arctic. Analyzing the movement data with a switching-state space model and a series of mixed effects models, we show that the presence of killer whales strongly alters the behavior and distribution of narwhal. When killer whales were present (within about 100 km), narwhal moved closer to shore, where they were presumably less vulnerable. Under predation threat, narwhal movement patterns were more likely to be transiting, whereas in the absence of threat, more likely resident. Effects extended beyond discrete predatory events and persisted steadily for 10 d, the duration that killer whales remained in Admiralty Inlet. Our findings have two key consequences. First, given current reductions in sea ice and increases in Arctic killer whale sightings, killer whales have the potential to reshape Arctic marine mammal distributions and behavior. Second and of more general importance, predators have the potential to strongly affect movement behavior of tracked marine animals. Understanding predator effects may be as or more important than relating movement behavior to resource distribution or bottom-up drivers traditionally included in analyses of marine animal tracking data.

Dietary habits of polar bears in Foxe Basin, Canada: possible evidence of a trophic regime shift mediated by a new top predator.

Polar bear (Ursus maritimus) subpopulations in several areas with seasonal sea ice regimes have shown declines in body condition, reproductive rates, or abundance as a result of declining sea ice habitat. In the Foxe Basin region of Nunavut, Canada, the size of the polar bear subpopulation has remained largely stable over the past 20 years, despite concurrent declines in sea ice habitat. We used fatty acid analysis to examine polar bear feeding habits in Foxe Basin and thus potentially identify ecological factors contributing to population stability. Adipose tissue samples were collected from 103 polar bears harvested during 2010-2012. Polar bear diet composition varied spatially within the region with ringed seal (Pusa hispida) comprising the primary prey in northern and southern Foxe Basin, whereas polar bears in Hudson Strait consumed equal proportions of ringed seal and harp seal (Pagophilus groenlandicus). Walrus (Odobenus rosmarus) consumption was highest in northern Foxe Basin, a trend driven by the ability of adult male bears to capture large-bodied prey. Importantly, bowhead whale (Balaena mysticetus) contributed to polar bear diets in all areas and all age and sex classes. Bowhead carcasses resulting from killer whale (Orcinus orca) predation and subsistence harvest potentially provide an important supplementary food source for polar bears during the ice-free period. Our results suggest that the increasing abundance of killer whales and bowhead whales in the region could be indirectly contributing to improved polar bear foraging success despite declining sea ice habitat. However, this indirect interaction between top predators may be temporary if continued sea ice declines eventually severely limit on-ice feeding opportunities for polar bears.

Prey items and predation behavior of killer whales (Orcinus orca) in Nunavut, Canada based on Inuit hunter interviews.

BACKGROUND: Killer whales (Orcinus orca) are the most widely distributed cetacean, occurring in all oceans worldwide, and within ocean regions different ecotypes are defined based on prey preferences. Prey items are largely unknown in the eastern Canadian Arctic and therefore we conducted a survey of Inuit Traditional Ecological Knowledge (TEK) to provide information on the feeding ecology of killer whales. We compiled Inuit observations on killer whales and their prey items via 105 semi-directed interviews conducted in 11 eastern Nunavut communities (Kivalliq and Qikiqtaaluk regions) from 2007-2010. RESULTS: Results detail local knowledge of killer whale prey items, hunting behaviour, prey responses, distribution of predation events, and prey capture techniques. Inuit TEK and published literature agree that killer whales at times eat only certain parts of prey, particularly of large whales, that attacks on large whales entail relatively small groups of killer whales, and that they hunt cooperatively. Inuit observations suggest that there is little prey specialization beyond marine mammals and there are no definitive observations of fish in the diet. Inuit hunters and elders also documented the use of sea ice and shallow water as prey refugia. CONCLUSIONS: By combining TEK and scientific approaches we provide a more holistic view of killer whale predation in the eastern Canadian Arctic relevant to management and policy. Continuing the long-term relationship between scientists and hunters will provide for successful knowledge integration and has resulted in considerable improvement in understanding of killer whale ecology relevant to management of prey species. Combining scientists and Inuit knowledge will assist in northerners adapting to the restructuring of the Arctic marine ecosystem associated with warming and loss of sea ice.

Loss of Arctic sea ice causing punctuated change in sightings of killer whales (Orcinus orca) over the past century.

Killer whales (Orcinus orca) are major predators that may reshape marine ecosystems via top-down forcing. Climate change models predict major reductions in sea ice with the subsequent expectation for readjustments of species' distribution and abundance. Here, we measure changes in killer whale distribution in the Hudson Bay region with decreasing sea ice as an example of global readjustments occurring with climate change. We summarize records of killer whales in Hudson Bay, Hudson Strait, and Foxe Basin in the eastern Canadian Arctic and relate them to an historical sea ice data set while accounting for spatial and temporal autocorrelation in the data. We find evidence for "choke points," where sea ice inhibits killer whale movement, thereby creating restrictions to their Arctic distribution. We hypothesize that a threshold exists in seasonal sea ice concentration within these choke points that results in pulses in advancements in distribution of an ice-avoiding predator. Hudson Strait appears to have been a significant sea ice choke point that opened up .approximately 50 years ago allowing for an initial punctuated appearance of killer whales followed by a gradual advancing distribution within the entire Hudson Bay region. Killer whale sightings have increased exponentially and are now reported in the Hudson Bay region every summer. We predict that other choke points will soon open up with continued sea ice melt producing punctuated predator-prey trophic cascades across the Arctic.

Phylogeny and divergence of the pinnipeds (Carnivora: Mammalia) assessed using a multigene dataset.

BACKGROUND: Phylogenetic comparative methods are often improved by complete phylogenies with meaningful branch lengths (e.g., divergence dates). This study presents a dated molecular supertree for all 34 world pinniped species derived from a weighted matrix representation with parsimony (MRP) supertree analysis of 50 gene trees, each determined under a maximum likelihood (ML) framework. Divergence times were determined by mapping the same sequence data (plus two additional genes) on to the supertree topology and calibrating the ML branch lengths against a range of fossil calibrations. We assessed the sensitivity of our supertree topology in two ways: 1) a second supertree with all mtDNA genes combined into a single source tree, and 2) likelihood-based supermatrix analyses. Divergence dates were also calculated using a Bayesian relaxed molecular clock with rate autocorrelation to test the sensitivity of our supertree results further. RESULTS: The resulting phylogenies all agreed broadly with recent molecular studies, in particular supporting the monophyly of Phocidae, Otariidae, and the two phocid subfamilies, as well as an Odobenidae + Otariidae sister relationship; areas of disagreement were limited to four more poorly supported regions. Neither the supertree nor supermatrix analyses supported the monophyly of the two traditional otariid subfamilies, supporting suggestions for the need for taxonomic revision in this group. Phocid relationships were similar to other recent studies and deeper branches were generally well-resolved. Halichoerus grypus was nested within a paraphyletic Pusa, although relationships within Phocina tend to be poorly supported. Divergence date estimates for the supertree were in good agreement with other studies and the available fossil record; however, the Bayesian relaxed molecular clock divergence date estimates were significantly older. CONCLUSION: Our results join other recent studies and highlight the need for a re-evaluation of pinniped taxonomy, especially as regards the subfamilial classification of otariids and the generic nomenclature of Phocina. Even with the recent publication of new sequence data, the available genetic sequence information for several species, particularly those in Arctocephalus, remains very limited, especially for nuclear markers. However, resolution of parts of the tree will probably remain difficult, even with additional data, due to apparent rapid radiations. Our study addresses the lack of a recent pinniped phylogeny that includes all species and robust divergence dates for all nodes, and will therefore prove indispensable to comparative and macroevolutionary studies of this group of carnivores.

How seals divide up the world: environment, life history, and conservation.

Pinnipeds display a remarkable variation in life history adaptations while successfully inhabiting almost every marine environment. We explore how they have done this by grouping the world's pinniped species according to their environmental conditions, mating systems, lactation strategies, and timing of life histories. Next, we tested whether any of these clusters provide information about risk of extinction (using the International Union for Nature and the Conservation of Natural Resources status ranks). Seals at risk were not characterized by differences in lactation pattern (22% short vs. 46% long), mating system (24% multi-male vs. 35% harems), or timing of life history events (23% fast vs. 42% slow) but did differ based on four environmental groupings. Grouping traits (rather than seals) described two clusters: one that included the environmental trait, primary productivity, and a second one that included all other environmental variables (seasonality, latitude, and temperature). Based on this result and theoretical considerations, we plotted seals according to energy (primary productivity) and variation (seasonality) and found a pattern analogous to that of the same four groups determined by cluster analysis of all environmental variables. Of the two pinniped groups representing low variation (equatorial and high productivity), ten of 21 seal species have been designated at risk, in contrast to none of the 13 seal species adapted to high variation. We conclude that seals appear to be best adapted to seasonal environments and thus, conservation efforts may benefit by concentrating on species inhabiting less variable environments.