Wolves, whales, and elephants: animal personalities, culture, and conservation

How studying animal culture can inform conservation.

“Mica, Maga’s daughter, patrols the shoreline with two-year-old Piqui, her youngest, and her five-year-old daughter, Aiken. As soon as Piqui ventures into water that is a little bit too shallow for her mom’s comfort, Mica positions herself quickly between her calf and the beach, guiding Piqui out towards deeper water. Piqui is still too young to be a strander, while Aiken, having reached an age when some precocious orcas become successful stranders, has been practicing the stranding technique continuously this season…today repeatedly retrieving a piece of driftwood from the beach.”

Valeria Vergara, Punta Norte, Argentina, 2019

“As she is foraging, Paw Ray lets out a long low rumble, barely within range of the human ear. Her two-year-old calf, Toh Gwah, usually stays close, but this morning his curiosity has prevailed, and he is down the hill out of sight. Paw Ray lets out another rumble. This time it’s much louder, some might say a bit more demanding. Little Toh Gwah emerges from the thick brush a hundred meters below. Encouraged by the sounds of her continuous rumbles, Toh Gwah eventually reaches mom. After they extend their trunks towards each other in greeting, Toh Gwah grabs a single blade of grass from Paw Ray’s mouth, sampling what could potentially be a new food item.”

Chelsea Greer, Thailand, 2017 

Although detailing a moment in the lives of two completely different species, these field notes have much in common. Both describe matriarchal, long-lived, cultural species. And both describe the behaviour of highly intelligent, socially complex individuals whom we got to know deeply during our respective field studies. 

Mica is a 19 year-old killer whale (Orcinus orca), daughter of 40-year old Maga, the matriarch of a family of 11, most of which are stranders. This means that they use a sophisticated and risky hunting strategy that involves charging at the shore, intentionally stranding themselves to seize Southern sea lion pups (Otaria flavescens), rolling back into the water as the surf recedes. Not surprisingly, learning and practicing to perfect this technique takes many years. Maga’s matriline is one of only three related orca matrilines that sustain this culturally transmitted behaviour in the waters of Peninsula Valdes, Patagonia, a UNESCO World Heritage site of global significance for the conservation of marine mammals.  

Paw Ray is a 29-year-old Asian elephant (Elephas maximus), the matriarch of a family recently reintroduced to their native habitat in northern Thailand. Although she spent a decade of her life giving rides to tourists, Paw Ray has acquired extensive social and ecological knowledge roaming the forest in her former 18 years. The selection and manipulation of a wide diversity of plants is an opportunity and a choice that is often deprived in captivity, which can be detrimental to the learning and development of critical foraging skills. Now, in their new forest home, Paw Ray can apply her vast memory and knowledge of the landscape to assist her young ones, like Toh Gwah, as they learn to socialize, forage, and navigate the complexities of their natural habitat.

A large killer whale stranding in shallow water while sea lion pups try to escape up the beach.
Male killer whale, Jaluel. Photo by Valeria Vergara.
A family of elephants walking in the forest.
Family of Asian elephants exploring their environment. Photo by Mahouts Elephant Foundation.

A paradigm shift: critical anthropomorphism

The field of biology has seen a gradual but steady shift in how scientists approach the animals they study, from viewing them as mere representatives of their species, to recognizing and valuing the individuality and complexity of each animal as a unique sentient being with her or his own personality, preferences, behaviours, and experiences. Kristy Ferraro and colleagues’ recent paper “Revisiting two dogmas of conservation science” beautifully encapsulates this shift.1 The authors argue that conservationists have traditionally ignored the value of individuals and have focused instead on the value of ecological collectives. They propose that a departure from the traditional focus on population, species, and ecosystem scales, and a consideration of individuals as the unit of inquiry can lead to a more comprehensive understanding of species and ecosystems and, ultimately, better conservation efforts.

Ferraro and her colleagues also challenge the conventional view that anthropomorphism should be avoided in conservation science and propose that critical anthropomorphism is a legitimate scientific tool. By relying on natural history, perceptions, intuitions, careful behaviour descriptions, and by “daring to imagine a shared experience”, critical anthropomorphism can lead to a more robust understanding of how animals interact with their environment and each other, and elicit a more compassionate approach to conservation.

As conservation biologists who have had the privilege to devote much of our lives to observing non-human animals, we are heartened by the growing recognition that individual differences matter, and that paying attention to the quirks and unique personalities of the animals we study is not only advisable, it is imperative. A more nuanced understanding of the particular characteristics and personality traits of individual animals catalyzes a deeper insight into the species as a whole. This approach can enrich our understanding of not only the complexities of animal behaviour, social dynamics, and cultural traditions, but also the effect of environmental changes and anthropogenic activities on specific populations. It is ultimately a more ethical approach to studying and interacting with our non-human counterparts, one that fosters their welfare and conservation

Tracking individuals visually

This shift has been made possible in part by advances in methods that make it easier to track and observe individual animals over time. Orca photo-identification studies are an excellent example. To anyone spending enough time with killer whales, their individually distinct saddle patches and eye patches can become as recognizable as human faces. Orca photo-identification studies in the Pacific Northwest, pioneered by Bigg and colleagues,2 have contributed to understanding much of their biology, including their life histories, social systems, movement patterns, site fidelity, and abundance.3 4 Even genetic, acoustic, and foraging studies have benefitted from photo-identification techniques.5 6 7

Killer whales at the surface of the water with Achiever in the background.
A23 matriline and Raincoast’s research sailboat, Achiever. Photo by Raincoast / Oceanwise.

Several other cetacean species can also be identified and tracked using similar methods. For example, humpback whales (Megaptera novaeangliae) can be distinguished by the unique patterns found on the underside and trailing edge of their tail flukes, much like human fingerprints that are unique to individuals. By photographing individual whales and their calves every year it becomes possible to establish family relationships, and to link particular individuals to their breeding and feeding grounds, and even to particular innovative behaviours, as we illustrate later through the story of Conger and Moonstar.  

A humpback whale tail sticks out of the water in front of a mountainous landscape.
Gandalf (BCZ0333), a humpback whale first sighted in 2014. Photo by Valeria Vergara / Raincoast Conservation Foundation.

In the terrestrial world, research is also progressing on photographic capture-recapture of megafaunal species using natural markings. Similar to the scars and notches that help identify killer whales (in addition to their unique saddle patches and eye patches), elephants also naturally acquire marks over their lifetime. Studies of African savannah elephants (Loxodonta africana) have largely used patterns of notches and tears on their large triangular ears to identify individuals.8 9 Vidya and colleagues found that for Asian elephants, in addition to ear features, tusk and tail characteristics, and body wounds and warts were most suitable trait states for reliable individual identification because of the variability across individuals and slower rate of change of those trait states.10

Research on terrestrial large carnivores has used distinctive features, such as unique facial markings (e.g., snow leopards, Panthera uncia),11 spot patterns (e.g., cheetahs, Acinonyx jubatus),12 and chest blazes (e.g., American black bears, Ursus americanus, and Asian black bears, Ursus thibetanus)13 14 to identify individuals and estimate density, annual survival, and demographic parameters of animals. Gray wolves (Canis lupus), on the other hand, can be more difficult to distinguish as their pelage can be subject to seasonal change. However, high quality photos can capture unique patterns and colours of facial markings and chest blazes, and the appearance of the saddle and shoulder area is quite variable among individuals. Furthermore, the scent gland spot on wolves’ tails can be helpful for differentiating individuals as its size, shape, and location remains consistent and readily visible throughout the year. For wildlife that lack distinctive markings, automated facial recognition has also proven to be a useful tool for non-invasively identifying individuals (e.g., BearID).15

A wolf walking in front of a camera trap.
Photo by Chelsea Greer / Raincoast Conservation Foundation.

Tracking individuals using sound

Acoustic monitoring using autonomous recording units (ARUs) are transforming the way in which we study individuals, groups, and populations of animals in both marine and terrestrial environments. Wolves exemplify this development beautifully. Much like us, every wolf has their own distinct voice, and as social beings, they use their voice (barks, growls, whimpers, howls) to communicate and bond with one another.16 17 By recording and analyzing wolf howls, which can travel distances up to several kilometers, researchers have been advancing a new way of identifying individual wolves.18 19 20 21 Holly Root-Gutteridge and colleagues used fundamental frequency and amplitude of howls to identify individual captive and wild Eastern gray wolves (C. l. lycaon).22 23 Moreover, researchers have noted howl dialects that correspond to specific subspecies of wolves,24 as well as specific packs within a subspecies.25 Coupled with camera traps, ARUs may improve long-distance detection of wolves and other canid species while also providing valuable insight into the environment in which these ecologically important predators live. 

Chorus wolf howl recorded as part of the Haíɫzaqv Wolf and Biodiversity Monitoring Project (Kyle Artelle/HIRMD).

Underwater, sound travels much more efficiently than light. Consequently, acoustic communication plays a crucial role in the behaviour and social interactions of marine creatures, particularly marine mammals.

Cetaceans rely on sound for nearly all aspects of their lives: to maintain social relationships, for group cohesion, as a signal of individual or group identity, to attract mates, to navigate, and to find food. Because of this, passive acoustic monitoring (PAM) can be an extremely effective non-intrusive method to not only study their habitat use, migration patterns, and foraging activity, but also to offer insights into their behaviour and social structure. PAM methods enable researchers to identify and track particular family units, pods, and clans, and even individuals by their sounds alone, particularly for well-studied species such as killer whales, sperm whales (Physeter macrocephalus), common bottlenose dolphins (Tursiops truncatus), and more recently beluga whales (Delphinapterus leucas) (a study Raincoast is currently involved with). 

A person standing on a tall platform while holding wires to set up a hydrophone.
Valeria setting up hydrophone to record beluga whales. Photo by Gretchen Freund.
Recording of A-Clan, one of the three Northern Resident Killer Whale acoustic clans (killer whale pods that share one or more calls belong to a common acoustic clan). Recording by Lance Barrett-Lennard.

Cultural species

Individual differences in behaviour are, of course, what drives evolution and genetic diversity. Individuals also play a vital role in maintaining and transmitting cultural traditions, and there is now ample evidence that such cultural traditions are not just limited to humans. Whales, wolves, and elephants are amongst the growing number of species known to possess unique cultural practices that are passed down from one generation to the next. 

What exactly do we mean by culture? Human culture tends to be defined as the way in which we do things, encompassing our values, beliefs, traditions, the way we dress, or the way we eat. But this definition of culture comes from cultural anthropology, which centers on criteria that apply only to humans. For the concept of culture to be useful when we study non-human animals, two fundamental criteria must be met: the transmission of information through social learning and the resulting shared behaviour exhibited by a group. In their seminal book “The Cultural Lives of Whales and Dolphins”, Hal Whitehead and Luke Rendell therefore adopted a definition of culture widely used by evolutionary biologists: culture is knowledge, information, or behaviour that is shared within a group or society of individuals and is acquired through some form of social learning.26

The power of personality

Animal individuality and distinct personalities can play a significant role in shaping culture within a group or society of animals. Bold individuals, for example, may be more likely to take risks and try new behaviours, while shy individuals may be more cautious and conservative in their actions. A bold and adventurous wolf might lead the pack to explore new territories, whereas a shy and cautious individual might prefer to remain in more familiar areas. These distinct personalities can influence the transmission of information within a group, leading to cultural traditions that can be group-specific. 

The importance of personality, especially of those in leadership roles, is exemplified in the story of 06, an alpha27 female wolf in Yellowstone National Park. 06 was a strong, intelligent, competent wolf who could take down large prey animals, such as elk (Cervus elaphus) by herself, which is extremely rare as wolves usually hunt in packs. When it came to raising her family, she was patient, cooperative, and steadfast. These characteristics were stark in contrast to the violent alpha female, 686F, from a neighboring rival pack. She was known to be excessively aggressive, and her pack killed at least nine other wolves under her tumultuous reign. These two female wolves were both leaders of their respective families, and their practices, whether it be cooperation or violence, may be passed down from generation to generation for as long as the pack persists

Personalities and individual preferences can also be important in group dynamics and ecosystem function. Increasingly, research emphasizes the importance of intraspecific variation and the disproportionate effect individuals can have on community structure and ecosystem function.28 For example, researchers in Voyageurs National Park have observed behaviours that indicate variation in wolf personalities, specifically in their hunting preferences and strategies. Some individuals hunted and ambushed beavers 229% and 263% more than other pack members, respectively.29 These findings show that some wolves in the Park have “beaver hunting personalities”, and interestingly, these personality-types can have a larger ecological effect, relative to the wolf population as a whole, because their hunting preferences and success rates may alter the formation of additional beaver-created wetlands in the ecosystem. Variation in wolf hunting personalities, in particular individual preferences for specific prey species (e.g. moose [Alces alces] over elk), has been observed in other regions, and may be a precursor for cultural development within these wolf packs.

The significance of individual animals shaping cultural traditions within animal societies is also demonstrated in the story of Granny, the matriarch of J pod, a Southern Resident killer whale family group in the north Pacific. Granny, formally known as J-2, was at least 90 years old when she died in 2016. Despite not having given birth to a calf in more than 50 years, Granny likely played a vital role in the pod’s survival through her extensive ecological knowledge. A research team that reviewed hundreds of hours of video footage found that during years of low Chinook salmon (Oncorhynchus tshawytscha) abundance, post-reproductive females, like Granny, were the ones leading the movements of the whales, as they possessed the greatest knowledge of where to find salmon. Grandmothers serve as the keepers of ecological knowledge within their family groups. As such, Granny was not interchangeable. Her individual personality and behaviour and her elderly wisdom contributed to shape the culture and traditions of her family group. A consequence of social learning can be the increased importance of key individuals as repositories of accumulated knowledge. From an applied conservation perspective, it would seem that targeting protection of these individuals would be of particular importance for the assuring persistence of social units.

Innovative individuals can also play an important role in introducing and disseminating feeding techniques in many species;humpback whales provide a wonderful example. These whales are generalist-feeders, using a range of strategies such as lunge-feeding and bubble-net feeding to hunt for food. In 2011, researchers at the Marine Education and Research Society (MERS), observed a novel hunting strategy off northeast Vancouver Island, British Columbia appropriately named “trap feeding”: the strategy involves the whale remaining almost stationary at the surface of the water with its mouth open for an extended period of time, resembling a Venus Flytrap. At first, only two individuals, adult male Conger, and three-year old Moonstar, were engaging in this behaviour. When Christie MacMillan, Jared Towers, and Jackie Hildering completed a study on trap feeding in 2015, 16 individuals had learned this behaviour since its discovery in 2011.30 And by 2018, more than 20 individuals were known to engage in this behaviour. Interestingly, none of the trap-feeders learned the behaviour from their mothers, but rather from one another, highlighting the importance of cultural transmission. 

Multiple humpback whales with only their mouths above water while bubble-net feeding.
Humpbacks bubble-net feeding. Photo by Valeria Vergara / Raincoast Conservation Foundation.

Lessons for conservation

The innovative humpback trap-feeding behaviour is thought to be linked to changes in ocean conditions, such as dwindling fish stocks, and it provides an energetically efficient method of feeding on small, diffuse prey patches. This underscores the idea that innovative individuals in animal societies can play an important role in introducing and disseminating behavioural techniques that can have significant conservation value.

Food preferences and foraging strategies are not the only forms of knowledge and practice that can be passed on through generations; so can trauma, which can negatively affect conservation outcomes. For elephants residing on both the African and Asian continents, poaching and live capture for captivity, respectively, pose major threats to their survival, welfare, and conservation. The loss of elders, maternal separation, and witnessing the killings of their family members impairs the normal development of young elephant brains and behaviour. Such trauma can be socially, culturally, and even neurobiologically transmitted across individuals and family groups.31 

In the article “Elephant breakdown,” trans-species psychologist Gay Bradshaw and her colleagues state that such trauma experienced by these animals can define a culture.32 What does this look like? Studies of various assaults and killings of rhinoceroses (Diceros bicornis, Ceratotherium simum) in South Africa have identified the perpetrators in all cases as young male elephants that had witnessed the mass killings of their families. Unprovoked elephant attacks on villages have also been reported in several African countries. Bradshaw and colleagues put forth the argument that present-day elephant populations are experiencing a type of persistent stress, akin to a collective trauma affecting the entire species. They contend that extensive periods of poaching, selective killing, and habitat destruction have profoundly disturbed the intricate network of familial and societal connections through which young elephants have historically been nurtured in their natural habitat. As a result, established elephant herds are facing what can be described as nothing short of a “​​precipitous collapse of elephant culture”, further exacerbating existing conflict risks between elephants and humans, and threatening the conservation of wildlife within those ecosystems. 

Side profile of elephant calf behind leafy vegetation.
Photo by Mahouts Elephant Foundation.

Why study animal cultures

“What’s at stake is: ways of being. What ancient memory banks are being purged, what rich files of life’s library are being erased? What’s at stake is communities of individuals who know who they are in the world because they know one another.”

Carl Safina, in Becoming Wild

As we’ve come to appreciate the importance of personalities, social learning, cultural transmission, and innovation in animal societies, it has become abundantly clear that in the face of human-induced rapid environmental change and habitat degradation we must focus on maximizing the survival and reproductive success of individuals, social groups, and populations.33 When we strive to understand the ways in which animals adapt and innovate in response to changing environments, we can better protect and preserve unique and valuable populations. In his book “Becoming Wild: How Animal Cultures Raise Families, Create Beauty, and Achieve Peace”, Carl Safina makes a compelling case for why it is important to study animal cultures. Biodiversity is not just a “gene-pool thing”, he writes, “skills, traditions, and dialects that animals have innovated and passed along culturally are crucial to helping many populations survive and perpetuate.” 

For conservationists in the Pacific Northwest of North America, there is no better example than the plight of the endangered Southern Resident killer whales. Killer whales are a quintessential illustration of a species that embodies cultural traits. Distinct cultural groups have specialized foraging techniques, vocal dialects, and unique social and behavioural patterns that are transmitted across generations. Southern Resident killer whales rely heavily on Chinook salmon as their primary food source. In her book chapter “A bond through salmon, language, and grandmothers”, Misty MacDuffee, Raincoast’s Wild Salmon Program Director, explains why this is so: in a nutshell, Chinook have a high fat content, they are available year-round within the whales’ foraging range, and their large size allows for sharing among family members.

The decline in Chinook salmon numbers and size is known to be one of the primary causes for the lack of recovery of this population, compounded by water pollution and vessel traffic. The Southern Resident killer whale’s dependence on a single prey source and their cultural reluctance to switch food sources is a fundamental piece of information that highlights the importance of understanding foraging culture in conservation management.

Equally important is understanding how a species’ evolutionary history influences its thinking process, social learning strategies, and group and population composition, which in turn affect how they may respond to human-induced rapid environmental change.34 Species with vertical or oblique cultures, whereby behaviour is learned from parents or other members of previous generations, such as killer whales, can be tremendously conservative and adhere to conformist traditions that can limit adaptive responses to rapid environmental change, whereas horizontal cultures, where transmission occurs readily between members of the same generation, can be very effective in quickly changing population behavior in adaptive ways.35

Social learning strategies that drive culture, therefore, can determine the responses to environmental changes that alter the availability of preferred or optimal prey.36 Wolves, like killer whales, are considered ecosystem generalists that can exploit a wide diversity of niches and eat a variety of foods. But while orcas at specific locations have specialized in very particular prey and these feeding traditions tend to be conservative, wolves are more flexible and can adapt more readily because of their ability to switch prey.37 Understanding the types of cultures that we are studying, the prevalent social learning mechanisms and the degree of conservatism within those cultures, helps us understand how likely individuals or groups would be to change tactics when facing environmental problems. 

The development of distinct cultures through social learning by groups of wild animals is greatly influenced by the ecosystems they inhabit. Degrading these ecosystems means these cultures will likely wither away. In addition to habitat protection, recognizing and safeguarding the rich cultural diversity of animal species is imperative to ensure their best chance for survival in the face of unprecedented environmental change and habitat degradation. 

Until recently, conservation strategies and policies have focused primarily on broad demographic responses and the preservation of genetically defined, evolutionarily significant units. Yet, animal culture has important consequences for the survival and reproduction of individuals, social groups, and potentially, entire populations. The notion that understanding animal culture is crucial for conservation has rightfully received considerable attention in recent years. By studying the intricate tapestry of animal culture and traditions, including personalities, innovative behaviours, distinct foraging techniques, communication systems, and social learning mechanisms, we not only honour their inherent value, but also gain invaluable knowledge to develop targeted and effective conservation strategies.

Acknowledgments

A special thanks to Dr. Paul Paquet for his insightful and illuminating contributions in the development and editing of this article.

Notes and references

  1. Ferraro, K. M., Ferraro, A. L., Arietta, A. Z. A., & Sommer, N. R. (2023). Revisiting two dogmas of conservation science. Conservation Biology. doi:10.1111/cobi.14101
  2. Bigg, M. A., Ellis, G. M., Ford, J. K. B. & Balcomb, K. C. (1987). Killer Whales: A Study of Their Identification, Genealogy, and Natural History in British Columbia and Washington State. Phantom Press and Publishers, Nanaimo, BC. 79 p.
  3. Ford, J. K. B. (2019). Killer Whales: Behavior, Social Organization, and Ecology of the Oceans’ Apex Predators. in Ethology and Behavioral Ecology of Odontocetes (ed. Würsig, B.) 239–259. Springer International Publishing. doi.org/10.1007/978-3-030-16663-2_11.
  4. Towers, J. R. (2020). Photo-identification catalogue and status of northern resident killer whale population in 2019. Canadian Technical Report of Fisheries and Aquatic Sciences 3371: iv + 69 p.
  5. Barrett-Lennard, L. G. (2000). Population structure and mating patterns of killer whales  (Orcinus orca) as revealed by DNA analysis. Doctoral dissertation. University of British Columbia.
  6. Yurk, H., Barrett-Lennard, L., Ford, J. K. B. & Matkin, C. O. (2002). Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska. Animal Behavior, 63, 1103–1119.
  7. Deecke, V. B., Barrett-Lennard, L. G., Spong, P. & Ford, J. K. B. (2010). The structure of stereotyped calls reflects kinship and social affiliation in resident killer whales (Orcinus orca). Naturwissenschaften, 97, 513–518.
  8. Bedetti, A., Greyling, C., Paul, B., Blondeau, J., Clark, A., Malin, H., Horne, J., Makukule, R., Wilmot, J., Eggeling, T., Kern, J., & Henley, M. (2020). System for elephant ear-pattern knowledge (SEEK) to identify individual African elephants. Pachyderm, 61, 63-77.
  9. Douglas-Hamilton, I. (1972). On the Ecology and Behaviour of the African Elephant. D. Phil. thesis, University of Oxford, Oxford, UK.
  10. Vidya, T. N. C., Prasad, D., & Ghosh, A. (2014). Individual identification in Asian elephants. Gajah, 40, 3-17.
  11. Alexander, J. S., Gopalaswamy, A. M., Shi, K., & Riordan, P. (2015). Face value: towards robust estimates of snow leopard densities. PLoS One, 10(8), e0134815.
  12. Chelysheva, E. V. (2004). A new approach to cheetah identification. Cat news, 41, 27-29.
  13. Reynolds-Hogland, M. J., Ramsey, A. B., Muench, C., Pilgrim, K. L., Engkjer, C., Erba, G., & Ramsey, P. W. (2022). Integrating video data with genetic data to estimate annual age-structured apparent survival of American black bears. Population Ecology, 64(4), 300-322.
  14. Ngoprasert, D., Reed, D.H., Steinmetz, R., & Gale, G.A. (2012). Density estimation of Asian bears using photographic capture–recapture sampling based on chest marks. Ursus, 23, 117–133.
  15. Clapham, M., Miller, E., Nguyen, M., & Darimont, C. T. (2020). Automated facial recognition for wildlife that lack unique markings: A deep learning approach for brown bears. Ecology and Evolution, 10(23), 12883-12892.
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  17. Joslin, P. W. B. (1967). Movements and home sites of timber wolves in Algonquin park. American Zoologist, 7(2), 279-288.
  18. Tooze, Z.J., Harrington, F.H., & Fentress, J.C. (1990). Individually distinct vocalizations in timber wolves, Canis lupus. Animal behaviour, 40(4), 723-730.
  19. Coscia, E.M., Phillips, D.P., & Fentress, J.C. (1991). Spectral analysis of neonatal wolf Canis lupus vocalizations. Bioacoustics, 3(4), 275-293.
  20. Passilongo, D., Buccianti, A., Dessi-Fulgheri, F., Gazzola, A., Zaccaroni, M. and Apollonio, M., 2010. The acoustic structure of wolf howls in some eastern Tuscany (central Italy) free ranging packs. Bioacoustics, 19(3), 159-175.
  21. Palacios, V., Font, E. & Márquez, R. (2007). Iberian wolf howls: acoustic structure, individual variation, and a comparison with North American populations. Journal of Mammalogy, 88(3), 606-613.
  22. Root-Gutteridge, H., Bencsik, M., Chebli, M., Gentle, L. K., Terrell-Nield, C., Bourit, A., Yarnell, R. W. (2013). Improving individual identification in captive Eastern Grey Wolves (Canis lupus lycaon) using the time course of howl amplitudes. Bioacoustics. doi.org/10.1080/09524622.2013.817318
  23. Root-Gutteridge, H., Bencsik, M., Chebli, M., Gentle, L. K., Terrell-Nield, C., Bourit, A., & Yarnell, R. W. (2014). Identifying individual wild Eastern grey wolves (Canis lupus lycaon) using fundamental frequency and amplitude of howls. Bioacoustics, 23(1), 55-66.
  24. Kershenbaum, A., Root-Gutteridge, H., Habib, B., Koler-Matznick, J., Mitchell, B., Palacios, V., & Waller, S. (2016). Disentangling canid howls across multiple species and subspecies: Structure in a complex communication channel. Behavioural processes, 124, 149-157.
  25. Passilongo, D., Buccianti, A., Dessi-Fulgheri, F., Gazzola, A., Zaccaroni, M., Apollonio, M. (2010). The acoustic structure of wolf howls in some eastern Tuscany (central Italy) free ranging packs. Bioacoustics, 19(3), 159-175.
  26. Whitehead, H. & Rendell, L. (2015). The cultural lives of whales and dolphins. The University of Chicago Press.
  27. We use the term “alpha”, as did the author of ‘The Alpha Female Wolf: The Fierce Legacy of Yellowstone’s 06’, although others might not.
  28. Des Roches, S., Post, D. M., Turley, N. E., Bailey, J. K., Hendry, A. P., Kinnison, M. T., Schweitzer, J. A., & Palkovacs, E. P. (2018). The ecological importance of intraspecific variation. Nature Ecology and Evolution, 2(1), 57-64.
  29. Bump, J., Gable, T., Johnson‐Bice, S., Homkes, A., Freund, D., Windels, S., & Chakrabarti, S. (2022). Predator personalities alter ecosystem services. Frontiers in Ecology and the Environment, 20(5), 275-277.
  30. McMillan, C. J., Towers, J. R., & Hildering, J. (2019). The innovation and diffusion of “trap-feeding,” a novel humpback whale foraging strategy. Marine Mammal Science, 35, 779–796.
  31. Rizzolo, J. B., & Bradshaw, G. A. (2016). Prevalence and Patterns of Complex PTSD in Asian Elephants (Elephas maximus). Asian Elephants in Culture and Nature, 291–297.
  32. Bradshaw, G. A., Schore, A. N., Brown, J. L., Poole, J. H., & Moss, C. J. (2005). Elephant breakdown. Nature, 433(7028), 807-807.
  33. Brakes, P., Carroll, E. L., Dall, S. R., Keith, S. A., McGregor, P. K., Mesnick, S. L., Noad, M. J., Rendell, L., Robbins, M. M., Rutz, C., Thornton, A., Whiten, A., Whiting, M. J., Aplin, L. M., Bearhop, S., Ciucci, P., Fishlock, V., Ford, J. K. B., Notarbartolo di Sciara, G., Simmonds, M. P., Spina, F., Wade, P. R., Whitehead, H., Williams, J. & Garland, E. C. (2021). A deepening understanding of animal culture suggests lessons for conservation. Proceedings of the Royal Society B, 288(1949), 20202718. doi.org/10.1098/rspb.2020.2718
  34. Barrett, B., Zepeda, E., Pollack, L., Munson, A., & Sih, A. (2019). Counter-culture: Does social learning help or hinder adaptive responses to human-induced rapid environmental change? Frontiers in Ecology and Evolution, 7. doi.org/10.3389/fevo.2019.00183
  35. Whitehead, H., Rendell, L., Osborne, R. W., & Würsig, B. (2004). Culture and conservation of non-humans with reference to whales and dolphins: review and new directions. Biological Conservation, 120(3), 427-437. doi.org/10.1016/j.biocon.2004.03.017
  36. Roffler, G. H., Eriksson, C. E., Allen, J. M., & Levi, T. (2023). Recovery of a marine keystone predator transforms terrestrial predator–prey dynamics. Proceedings of the National Academy of Sciences, 120(5), e2209037120.
  37. Prokopenko, C. M. (2022). Hungry wolves and dangerous prey: a tale of prey switching (Doctoral dissertation, Memorial University of Newfoundland).

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Coastal wolf with a salmon in its month.
Photo by Dene Rossouw.