OUR RESEARCH
Center for Whale Research Research Director Dr. Michael Weiss (left) and ORCA SURVEY Lead Dave Ellifrit aboard our new research vessel, KCB III. Read about KCB III in the March/Spring 2024 of the WHALE Report, CWR’s quarterly member newsletter.
Every year for five decades, we have collected detailed demographic data on the Southern Resident killer whale population, recording all observed births and deaths.
ORCA SURVEY
Since 1976
Southern Resident killer whales (orcas)
Center for Whale Research staff has also gathered detailed information on the behavior and ecology of the Southern Resident killer whales (SRKW), including where they are in geographic location and time, and their social behavior and foraging patterns. Our dataset has provided ground-breaking insight into killer whale biology and ecology that we hope will help to inform management decisions to conserve this vulnerable and now endangered population.
Left-side ID photograph of SRKW male, L88, from 2023 OS Encounter #45 on August 19.
(Photograph by ORCA SURVEY Lead Dave Ellifrit).
Much of our recent research has focused on how social factors influence killer whale survival, reproduction, and health. Our photo-ID study has demonstrated how killer whale survival is dependent on the presence of mothers (Foster et al. 2012), grandmothers (Nattrass et al. 2019), and strong social ties (Ellis et al. 2017). We’ve also found that raising sons is costly for mothers, significantly reducing their reproductive success (Weiss et al. 2023). Finally, by analyzing thousands of images, we’ve shown how older females protect their sons from receiving injuries from other whales (Grimes et al. 2023).
Population Biology and Conservation
Our first encounter with the Southern Resident killer whales was on April 8, 1976, near Port Angeles in the Strait of Juan de Fuca. At this time very little was known about the population size and structure.
During the preceding two decades marine parks had taken (or killed during capture) over 50 individuals from the population (Asper and Cornell 1977). Published reports suggested an abundance of animals and that individuals could continue to be sustainably taken from the population (Asper and Cornell 1977). In sharp contrast, using photographic ID techniques that were pioneered by Dr. Michael Bigg, we showed that this was a very small and vulnerable population. In 1976 we agreed with Dr. Bigg that there were approximately 68 animals remaining (Balcomb and Goebel 1976), with approximately 40% of the population having been taken into captivity or killed during capture. This research helped to end the live captures of killer whales from the Pacific Northwest. Both through our long-term monitoring of the population and through molecular work (e.g., Ford et al. 2011 and 2017) we have established that the population is demographically closed (there is no emigration or dispersal from the population and no gene flow with neighboring populations).
The detailed information collected by the Center for Whale Research on the population dynamics of the Southern Resident killer whales has been used to support management decisions in both Canada and the United States. In 2001, the Southern Resident killer whales were listed as endangered in Canada under the Species at Risk Act and, in 2005, they were designated endangered in the USA under the federal Endangered Species Act. Currently, the Southern Resident killer whales are one of the most critically endangered populations of marine mammals in the USA (NMFS 2017) and Canada (DFO, 2017).
Through our annual monitoring of the population, we have been able to track the changes in population size, which has fluctuated considerably over the last five decades (Krahn et al. 2002; Olesiuk et al. 2005; Balcomb et al. 1982). From a starting point of just 68 identified individuals in 1976 following the live captures (Balcomb et al. 1982), the population showed signs of recovery and grew to a peak size of 98 individuals by 1995. The population then went into a steep decline with only 78 individuals remaining in the population by July 2000. This decline was followed by a further period of moderate growth to 89 individuals in the population in July 2006. Since then the population has declined once again, and as of December 2023, there are only 75 individuals remaining in the population, which is getting very close to the population size following the depletion of the population by the live captures. The population is not recovering.
Our long-term monitoring of this population has provided key insights into the cause of this decline. In particular, over the years we have contributed to studies to quantify the foraging behavior of the Southern Resident killer whales (Heimlich-Boran 1986; Ford et al. 1998; Ford and Ellis 2005). This work has established that Chinook salmon (Oncorhynchus tshawytscha) is the predominant prey species in the diet of the Southern Resident killer whale population during the summer months (Ford et al. 1998; Ford and
J3, J10, and other J pod whales in 1976
(Photograph by Ken Balcomb).
Ellis 2005) and that this dietary specialization is a major factor determining both their year-round distribution (Ford et al. 1998; Ford and Ellis 2005) and local habitat use (Heimlich-Boran 1986). This work has shown that periods of population decline (high mortality rates) are tightly associated with changes in Chinook salmon abundance, with higher mortality observed when salmon abundance is low (Ford et al. 1998; Ford and Ellis 2005). Moreover, our long-term data has shown that killer whale fecundity is highly correlated with the abundance of Chinook salmon with the probability of a female having a calf being 50% greater in years of high salmon abundance compared to years of low salmon abundance (Ward et al. 2009). These results highlight that the coast wide decline in Chinook abundance is likely to be the most significant factor contributing to the decline in the population.
In 2015 we provided written evidence for the hearing of the Canadian Trans Mountain Pipeline Expansion Project. Working with a team of international colleagues, we used population viability analysis (PVA) to assess the viability of the Southern Resident killer whale population and the impact that the proposed Trans Mountain Pipeline Expansion project would have on the population. This analysis showed that increased traffic and noise conditions would intensify existing threats, probably accelerating the rate of population decrease, possibly leading to complete loss of the population (Lacy et al. 2016).
It is often difficult to collect detailed information on individual animals in the marine environment, and we have worked with colleagues to develop new methods for collecting essential data to inform conservation and management decisions. These projects include using low power lasers to collect precise data on morphometrics and measure changes in the size and shape of the whales (Durban and Parsons 2006), and aerial images taken from a helicopter (Fearnbach et al. 2011. and 2017 in prep) to determine whale size, condition, and long-term growth trends. This data has been used to track the body condition and health of the whales over time. We are continuing to look for new ways of collecting data that will help both our understanding of these animals and inform their conservation and management. We now use an unmanned aerial vehicle (drone) to collect high-resolution video on the behavior of the whales (Aerial Observation Study).
Social Organization
Because all individuals in the population are individually identifiable due to their unique markings, our long-term monitoring of the population over the last five decades has provided detailed data on the population social structure.
We have established that neither sons nor daughters disperse from their mother and the population shows a stable matrifocal society composed of a hierarchy of social units. These matrilineal groups form three distinct pods (J, K, and L pods) that have persisted throughout the last five decades (Bigg 1990; Parsons et al. 2009; Foster et al. 2012b).
Although the overall pod structure has remained stable over decades, we have shown that the connectivity of the population has fluctuated and is also tightly linked to salmon abundance (Parsons et al. 2009; Foster et al. 2012b). Social groups are smaller in years of low salmon abundance (Parsons et al. 2009), and the population is less socially connected in years of low salmon (Foster et al. 2012b). Given the central importance of the social network for population processes such as mating dynamics and disease transmission, a change in social network structure caused by a change in food availability may have significant consequences for long-term population viability.
K45 and mother, K16, during 2024 OS Encounter #10 on January 30 (Photograph by Brendon Bissonnette).
By integrating our long-term data on the social structure of the population with molecular studies, we have been able to determine the mating dynamics in the population (Ford et al. 2011, and 2017 in review). This work has found high variance in male reproductive success with older and larger males having higher reproductive success. Although mating in the population often occurs in the same pod, current evidence suggests that mating between close relatives is avoided (Ford et al. 2017, in review) with rare exceptions.
The Evolution of Post Reproductive Females
One striking observation that emerged from our long-term studies is that much like humans, female resident killer whales typically have their last offspring in their late 30s or early 40s, but may live into their 80s and beyond (Olesiuk et al. 1990).
This menopausal life history is very rare; indeed the only other mammals reported to exhibit this unusual life history strategy are humans and short-finned pilot whales (Croft et al. 2015). The question of why females of any species should stop reproduction before the end of life is a long-standing evolutionary puzzle. One hypothesis is that post-reproductive females play a key role in helping their relatives to survive and reproduce. Mathematical models suggest that in killer whales post-reproductive females should invest more time and energy to support their sons rather than their daughters—in this way grand offspring from sons (who mate with a female from a different social group) will not compete with the family group, and post-reproductive females can maximize the number of grand offspring that they have (Johnstone and Cant 2010). We tested this hypothesis and found that post-reproductive females provide significant survival benefits to both sons and daughters, but these effects are much more pronounced in sons. If a post-reproductive female dies, the risk of her adult son dying in the year following her death is up to 8 times greater. In contrast, their adult female offspring only have a 2.7 fold increased risk (Foster et al. 2012a).
Our long-term observations of the population have shown that one way that females boost the survival of their group is by acting as repositories for ecological knowledge (Brent et al. 2015). As outlined above, Southern Resident killer whales feed almost exclusively on salmon and move into the inshore waters to feed on the salmon as they migrate back to their natal streams to spawn. The timing here is everything—the whales need to know when and where the salmon will be. By observing patterns of group travel and analyzing over 750 hours of video we found that it is the experienced post-reproductive females that guide their group around the salmon foraging grounds and that they especially do this in times of low salmon (Brent et al. 2015). However, we know that reproductive females can also act as leaders and support their adult sons.
Why then do the females stop reproducing?
Our research has shown that it is a combination of the help that older females can provide the group and the costs of reproductive competition caused by reproducing at the same time as their daughters that drives the evolution of menopause in Southern Resident killer whales. These are the very same mechanisms that are hypothesized to drive the evolution of menopause in humans.
Estimated 50-year-old J16 and her eldest son, J26, during 2023 OS Encounter #67 in Rosario Strait, Washington on October 18, 2023 (Photograph by ORCA SURVEY Lead Dave Ellifrit).
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(2017) Fisheries and Oceans Canada. Action Plan for the Northern and Southern Resident Killer Whale (Orcinus orca) in Canada. Species at Risk Act Action Plan Series. In, Fisheries and Oceans Canada, Ottawa
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Asper E, Cornell L (1977) Live capture statistics for the killer whale (Orcinus orca) 1961-1976 in California, Washington and British Columbia. Aquatic Mammals 5:21-26
-
Balcomb KC, Boran JR, Heimlich SL ( 1982.) Killer whales in greater Puget Sound. Reports of the International Whaling Commission 32: 681–685
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Bigg MA (1990) Social organization and genealogy of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. In: Hammond P, Mizroch S, Donovan G (eds), pp 383-405
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Brent LJN, Franks DW, Foster EA, Balcomb KC, Cant MA, Croft DP (2015) Ecological Knowledge, Leadership, and the Evolution of Menopause in Killer Whales. Current Biology 25:746-750
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Croft DP, Brent LJN, Franks DW, Cant MA (2015) The evolution of prolonged life after reproduction. Trends in Ecology and Evolution 30:407-416
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Croft DP, Johnstone Ra, Ellis S, Nattrass S, Franks DW, Brent LJN, Mazzi S, Balcomb KC, Ford JKB, Cant MA (2017) Reproductive Conflict and the Evolution of Menopause in Killer Whales. Current Biology 27:298-304
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Durban JW, Parsons KM (2006) Laser-metrics of free-ranging killer whales. Marine Mammal Science 22:732-743
-
Fearnbach H, Durban JW, Ellifrit DK, Balcomb KC (2011) Size and long-term growth trends of endangered fish-eating killer whales. Endangered Species Research 13:173-180
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Ford JKB, Ellis GM (2005) Prey selection and food sharing by fish-eating ‘resident’ killer whales (Orcinus orca) in British Columbia. In:
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Ford JKB, Ellis GM, Barrett-Lennard LG, Morton AB, Palm RS, Balcomb Iii KC (1998) Dietary specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters. Canadian Journal of Zoology 76:1456-1471
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Ford MJ, Hanson MB, Hempelmann JA, Ayres KL, Emmons CK, Schorr GS, Baird RW, Balcomb KC, Wasser SK, Parsons KM, Balcomb-Bartok K (2011) Inferred Paternity and Male Reproductive Success in a Killer Whale (Orcinus orca) Population. Journal of Heredity 102:537-553
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Foster Ea, Franks DW, Mazzi S, Darden SK, Balcomb KC, Ford JKB, Croft DP (2012a) Adaptive Prolonged Postreproductive Life Span in Killer Whales. Science 337:1313-1313
-
Foster Ea, Franks DW, Mazzi S, Darden SK, Balcomb KC, Ford JKB, Croft DP (2012a) Adaptive Prolonged Postreproductive Life Span in Killer Whales. Science 337:1313-1313
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Foster Ea, Franks DW, Morrell LJ, Balcomb KC, Parsons KM, van Ginneken A, Croft DP (2012b) Social network correlates of food availability in an endangered population of killer whales, Orcinus orca. Animal Behaviour 83:731-736
-
Heimlich-Boran JR (1986) Behavioral ecology of killer whales (Orcinus orca) in the Pacific Northwest. Canadian Journal of Zoology 66:565-578
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Johnstone RA, Cant MA (2010) The evolution of menopause in cetaceans and humans: the role of demography. Proceedings of the Royal Society B-Biological Sciences 277:3765-3771
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Krahn MM, Wade PR, Kalinowski ST, Dahlheim ME, Taylor BL, Hanson MB, Ylitalo GM, Angliss RP, Stein JE, Waples RS (2002) Status Review of Southern Resident Killer Whales (Orcinus orca) under the Endangered Species Act. In:
-
Lacy RC, Balcomb Iii KC, Brent LJN, Croft DP, Clark CW, Paquet PC (2016) Report on Population Viability Analysis model investigations of threats to the Southern Resident Killer Whale population from Trans-Mountain Expansion Project. In:
-
Olesiuk P, Bigg M, Ellis G (1990) Life history and population dynamics of resident killer whales (Ornicus orca) in the coastal waters of British Columbia and Washington State. Rep Int Whal Commn 12:209 - 243
-
Olesiuk PF, Bigg Ma, Ellis GM (2005) Life history and population dynamics of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State.209-243
-
Parsons KM, Balcomb KC, Ford JKB, Durban JW (2009) The social dynamics of southern resident killer whales and conservation implications for this endangered population. Animal Behaviour 77:963-971
-
Reynolds III JE, Marsh H, Ragen TJ (2009) Marine mammal conservation. Endangered Species Research 7:23-28
-
Ward EJ, Holmes EE, Balcomb KC (2009) Quantifying the effects of prey abundance on killer whale reproduction. Journal of Applied Ecology 46