The behavioral specializations, adaptations, energetics, and social patterns of PCFG gray whales

Dr. Clara Bird, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, and Conservation Sciences, GEMM Lab & LABIRINTO

In one of my first GEMM lab blogs (over six years ago!) I wrote that for my thesis I was going to, “…use the drone footage to analyze gray whale behavior and how it varies across space, time, and individual.”, and I’m happy to say that I more or less accomplished that goal.  Now as I write my last blog for the GEMM lab, a whole PhD and postdoc later, I want to take this opportunity to share what we’ve learned about Pacific Coast Feeding Group (PCFG) gray whale behavior from my PhD and postdoc work.

A behavioral specialization

Given the impressive diversity of foraging tactics used by PCFG gray whales (Torres et al., 2018), a central question from the start was, “do all individuals use all behaviors, or is there variation in which whales use each behavior?”. This interest in individual specialization led to several blogs and became the question I asked in my first PhD chapter (read an introduction to specialization here and summaries of the drivers of specialization here and here). In my first chapter, I used drone data to study the relationship between individual behavior use, body length and condition, and habitat type. We found a strong relationship between foraging behavior and individual length (which is also a proxy for age). Longer, older, whales were more likely to feed using the headstanding tactic while shorter, younger, whales were more likely to feed using forward swimming tactics (Figure 1; Bird et al., 2024a). Together, these results suggest an ontogenetic shift (i.e., a shift associated with age) in foraging behavior use. Furthermore, we found that different tactics were more likely to be used in different habitats; headstanding was more likely to occur in reef habitats while the forward swimming tactics were more likely to occur in rock habitat. Overall, this chapter showed us that PCFG gray whale foraging behavior varies by length/age and habitat, indicating a lack of generalization across the group.

Figure 1. The relationship between individual total length and the probability of a behavior being used. In each box, the x-axis represents total length, and the y-axis represents the probability of that behavior (shown in the box title) being used. Figure from Bird et al. (2024a).

A behavioral adaptation

If you’ve ever watched gray whales off the coast and seen a large bubble rise to the surface, then you’ve seen a bubble blast! While we observed these bubble blasts, described as “underwater release of air that rises to surface and forms a circle/puka.” (Torres et al., 2018), fairly often in the field (Figure 2), we were never quite sure of their function, leading to my second chapter.

Figure 2. Sequential photos extracted from drone video of bubble blasts performed by PCFG gray whales during a headstand (a), side-swim stationary (b), and subsurface feeding (c). Images 1–5 in each panel show a bubble blast event from the start of the exhalation (1) to the whale continuing to feed after the bubble has diffused at the surface (5). Figure from Bird et al., (2024b).

We initially wondered if bubble blasts served a prey corralling function (like humpback whale bubble nets), but the timing and location did not fit that idea. We instead wondered if bubble blasts were being used to regulate buoyancy. The whales we study forage in water nearly as shallow (<15 m) as they are long (~12 m), meaning that they must work against their buoyancy to dive. So, like a diver releasing air from their vest to sink, we hypothesized that these whales release air from their lungs (in the form of a bubble blast) to be able to dive more efficiently. Building on this idea, we specifically hypothesized that a whale would be more likely to bubble blast if they were bigger (i.e., because they had larger lungs) and fatter (i.e., they are more buoyant due to increased blubber). To test this hypothesis, we modeled the relationship between bubble blast use, total length, and body condition and found that the probability of an individual whale bubble blasting increased with total length and body condition. Furthermore, we found that whales who bubble blasted performed longer dives than those who did not, supporting our hypothesis that bubble blasts improved dive efficiency (Bird et al., 2024b).

Behavior and energetics

The interpretation of results from my first two chapters involved many questions regarding energetics. As we’ve described in previous blogs (here and here), it is important to understand how much energy different behaviors require because energetics helps us understand foraging success. Following the results of my first chapters, we wanted to better understand if different foraging behaviors cost different amounts of energy and if bubble blasts affected the energetic cost of a dive. To ask these questions we used individual breathing patterns as a proxy for energy expenditure (read more on the method here) and explored how breathing patterns were related to individual length, body condition, and behavior (including dive duration, foraging tactic, and bubble blast use). We found that the energetic cost of a dive increased with individual length, body condition, and dive duration (Figure 3.A1-3). Interestingly, we found no relationship between foraging tactic, bubble blast use and energetic expenditure (Bird et al., 2025; Figure 3.A4). However, my second chapter showed that both foraging behavior and bubble blast use affect dive duration (Bird et al., 2024b), indicating that effects of behavior on energetics come via the dive duration variable.

Figure 3. Estimated relationships between (1) total length (TL), (2) Body Area Index (BAI), (3) preceding dive duration (s) and (4) preceding dive foraging tactic and bubble blast occurrence and (A) total inhalation duration (s). Here total inhalation duration is the sum of all inhalations following a dive; a higher value indicates higher energy expenditure during the dive. In A4 the foraging tactics have been abbreviated as follows: HS = Headstand, Side.Sw.St = Side-swim stationary, Fwd.Sw. = Forward swimming tactics, Sub.St = Subsurface stationary, Surf. = Surface tactics. Figure from Bird et al., (2025).

Social patterns

As a postdoctoral scholar I had the opportunity to pivot from PCFG foraging behavior to social behavior. We generally think of baleen whales as solitary animals with loose social structure when on their foraging grounds, including gray whales while in nearshore Oregon waters. But social structure is not well studied in gray whales and can provide important insight into how information or disease might pass through a population. To look for social patterns we first assigned whales to a group if they were seen within 10 minutes and 100 meters of each other; whales seen in the same group were determined to be “associated”. If we saw whales interact with each other (e.g., touch each other, swim in a synchronized movement) they were determined to be “interacting”. We then tallied the number of times each possible pair of whales had been seen associating and/or interacting. The higher the tally, the stronger the association. Using that dataset, we assessed if some whales were more central (i.e., had strong associations or more associations with other whales) than others and if centrality was related to sex and age. We also assessed if whales were more likely to associate with other whales of similar sex or age. Finally, we reviewed our notes from the field and drone footage and documented the kinds of social interactions we’ve observed. While we’re still wrapping up this work, I’m excited to share that we’ve found that gray whales have more social structure than previously thought, including relationships with age and sex, and documented several interesting social interactions (Figure 4). I am excited to see what more years of data collection reveal about their social patterns, especially with an emphasis on how they might be learning from each other.

Figure 4. A social interaction documented from the drone. Here one whale is pursuing the other. Collected under NMFS permit #27426.

Tying it all together

Looking ahead, I’m most curious to better understand how the PCFG successfully feed in this shallow habitat. The findings of my third chapter show that the energetic cost of foraging increases with body condition (Bird et al., 2025). I hypothesize that this increase is because it becomes physically more difficult to dive as they become more buoyant (due to the increased fat). So, while bubble blasts appear to be a behavioral adaptation to reduce buoyancy (Bird et al., 2024b), there could be a point at which a whale is too fat to continue feeding in this shallow environment. Could this be why PCFG gray whales are skinnier than the Eastern North Pacific (ENP) gray whales that feed in the deeper arctic waters (Torres et al., 2022)? Given recent evidence that the PCFG may be facing a possible population decline (Pirotta et al., 2025), these questions are more relevant than ever.

The one theme that weaves throughout all this work is the importance of individual variation. Thanks to our incredible dataset, built from years of hard work and accessible whales that keep returning to our study site, we are able to follow individuals over time and uncover the links between habitat, individual size, body condition and sex, behavior, energetics, and the whales themselves. 

While I am sad to be leaving the GEMM lab, I am certainly proud of all that we have learned so far and excited to see what’s next (as an avid reader of the blog of course).

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References

Bird CN, Pirotta E, New L, Bierlich KC, Donnelly M, Hildebrand L, Fernandez Ajó A, Torres LG. 2024a. Growing into it: evidence of an ontogenetic shift in grey whale use of foraging tactics. Animal Behaviour 214:121–135. DOI: 10.1016/j.anbehav.2024.06.004.

Bird CN, Pirotta E, New L, Bierlich KC, Hildebrand L, Fernandez Ajó A, Torres LG. 2024b. Bubble blasts! An adaptation for buoyancy regulation in shallow foraging gray whales. Ecology and Evolution 14:e70093. DOI: 10.1002/ece3.70093.

Bird CN, Pirotta E, New L, Cornelius JM, Sumich JL, Colson KM, Bierlich KC, Hildebrand L, Ajó AAF, Doron A, Torres LG. 2025. Size and body condition drive the energetic cost of a baleen whale foraging in shallow habitat. PeerJ13:e20247. DOI: 10.7717/peerj.20247.

Pirotta E, New L, Fernandez Ajó A, Bierlich KC, Bird CN, Buck CL, Hildebrand L, Hunt KE, Calambokidis J, Torres LG. 2025. Body size, nutritional state and endocrine state are associated with calving probability in a long-lived marine species. Journal of Animal Ecology 94:1–13. DOI: 10.1111/1365-2656.70068.

Torres LG, Bird CN, Rodríguez-González F, Christiansen F, Bejder L, Lemos L, Urban R J, Swartz S, Willoughby A, Hewitt J, Bierlich KC. 2022. Range-Wide Comparison of Gray Whale Body Condition Reveals Contrasting Sub-Population Health Characteristics and Vulnerability to Environmental Change. Frontiers in Marine Science 9:1–13. DOI: https://doi.org/10.3389/fmars.2022.867258.

Torres LG, Nieukirk SL, Lemos L, Chandler TE. 2018. Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science 5:1–14. DOI: 10.3389/fmars.2018.00319.

How Humans and Cetaceans Shape Each Other

Marc Rams i Rios, PhD Student, Oregon State University Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

When I moved to Oregon to begin my PhD, I pictured long days on the water watching gray whales feed and travel along the coast. That does happen, and it is as incredible as I imagined. But I have learned that studying cetaceans is about much more than observing whales. It is also about people: how cultures – past and present – perceive these animals and share space with them.

In addition to marine mammals, I have always loved history and geography. Now, as I start my work with the GRANITE Project in the GEMM Lab, I find myself thinking about how these relationships between humans and whales unfold across time and space. In this post, I want to share a few examples of how whales have shaped human traditions for hundreds, even thousands of years, across societies that have never crossed. Then I will discuss how our research fits into this larger picture of human–cetacean connections.

Our journey begins in India, where the Ganges River dolphin inhabits a river that millions of people consider sacred. Its presence has long been linked to the health of the river, giving the species spiritual and cultural significance. Over the past century, the river’s ecological integrity has declined due to pollution, altered flow, and habitat disturbances, and this has caused the dolphin population to diminish1, 2. Conservation efforts that improve water quality, restore natural flow, and reduce disturbances not only help the dolphin recover but also protect the river and the human communities that rely on it1, 2. In this way, cultural reverence for the dolphin drives conservation measures that benefit both people and ecosystems1, 2.

© WWF Mohd Shahnawaz Khan

From there we move to Aotearoa, New Zealand, where Māori tradition speaks of tohorā, or whales, as guardians and ancestors3. They appear in ancestral stories as guides and protectors, and whale strandings have historically brought communities together in collective response. The Māori principles of kaitiakitanga, or guardianship, continue to shape marine conservation decisions today, guiding policies that integrate ecological and cultural values4. Here, whales are not seen as resources. They are part of a living genealogy that binds people to the sea and the life it sustains. In fact, team members of the SAPPHIRE project in the GEMM lab frequently engage with multiple iwi (Māori tribes) across Aotearoa through hui (meetings) where knowledge, stories, and culture are shared about blue whales and their ecosystem.

Traveling nearly to the antipodes, we arrive on the Atlantic coast of Brazil, in the town of Laguna, where an extraordinary partnership has endured for centuries. Artisanal fishers work alongside bottlenose dolphins, who drive schools of fish toward the shore and signal the right moment to cast the nets5, 6, 7. This cooperation benefits both species, and the knowledge behind it is passed down through generations of humans and dolphins through observation and shared practice5, 6, 7. It is a powerful example of how species can learn from one another, creating connections that challenge the idea of humans and wildlife as competitors and showing the potential for collaboration across species5, 6, 7. The LABIRINTO Lab in MMI has studied this interspecific relationship for decades, helping us learn about the patterns and endurance of these cultures.

PELD-SELA: Long-term ecological project on the Laguna Estuarine System and Adjacent Areas Projects. (n.d.). https://thelabirinto.com/projects1/

At the top of the Americas, in the Arctic, Inuit communities have hunted bowhead whales for thousands of years. These hunts are not only a source of food but also form the foundation of cultural identity and social life8. Knowledge of the ice, weather, and whale behavior is passed down through generations, and the hunt itself is embedded in ceremonies and practices that sustain the community8. Today, these traditions continue under strict quotas set through international agreements, carefully balancing cultural continuity with conservation9. The MMBEL lab in MMI studies the communication and ecology of bowhead whales to support the survival of this iconic species and the culture of Inuit people.

Emory Kristoff, National Geographic

Finally, our journey brings us to Oregon, where gray whales feed along a coastline rich with reefs, kelp beds, and sandy bottoms. These waters support a variety of human activities, from commercial fishing to recreation, creating risks such as entanglement, vessel strikes, and disturbance10, 11. Even well-intentioned actions like whale watching can cause harm if not carefully managed12, 13. Around the world, many communities have shifted from whaling to whale watching, transforming former hunting grounds into tourism destinations. While this is a positive change, it still requires monitoring. Noise can stress whales, boats can disrupt their behavior, and too much interaction can alter natural feeding and social patterns12, 13. In Oregon, research on gray whale habitat use and feeding home ranges helps inform management and conservation14.

Tradewind Charters Whale Watching and Fishing

This is where project GRANITE, Gray whale Response to Ambient Noise Informed by Technology and Ecology, comes in15. The project studies how whales respond to human activities by using drones to monitor health and behavior, photo-ID to track individuals, prey mapping to understand feeding choices, and acoustic recorders to capture the soundscape15, 16, 17. Equally important is collaborating directly with fishers and resource managers to reduce risks and develop solutions that benefit both whales and people. Healthy whale populations support communities too, through ecotourism, cultural continuity, education, and the ecological services whales provide. Conservation is reciprocal: caring for whales strengthens the ocean systems that sustain us all.

The tools and techniques developed by GRANITE, including drones, acoustic monitoring, and prey mapping, are not limited to Oregon. They can be applied globally, contributing to the protection of cetaceans in diverse habitats15. In this way, Oregon becomes more than the final stop on our tour. It is a place where centuries of human–whale relationships, lessons from around the world, and modern science converge. These examples across the world remind us that conservation is about more than preventing harm. It is about fostering a future where humans and whales thrive together, as they have shared the ocean for millennia.

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References

1 Sinha, R. K., & Kannan, K. (2014). Ganges river dolphin: An overview of biology, ecology, and conservation status in India. AMBIO, 43(8), 1029–1046. https://doi.org/10.1007/s13280-014-0534-7

2 Braulik, G., Atkore, V., Khan, M. S., & Malla, S. (2021). Review of scientific knowledge of the Ganges river dolphin. WWF. https://riverdolphins.org/wp-content/uploads/2021/07/Ganges-River-dolphin-Scientific-Knowledge-Review-July2021.pdf

3 Taonga, N. Z. M. for C. and H. T. M. (n.d.). Whales in Māori tradition. Teara.govt.nz. https://teara.govt.nz/en/te-whanau-puha-whales/page-1

4 McAllister, T., Hikuroa, D., & Macinnis‑Ng, C. (2023). Connecting science to Indigenous knowledge: Kaitiakitanga, conservation, and resource management. New Zealand Journal of Ecology, 47(1), 3521. https://doi.org/10.20417/nzjecol.47.3521

5 Simões‑Lopes, P. C., Fabián, M. E., & Menegheti, J. O. (1998). Dolphin interactions with the mullet artisanal fishing on southern Brazil: A qualitative and quantitative approach. Revista Brasileira de Zoologia, 15(3), 709–726. https://doi.org/10.1590/S0101-81751998000300008

6 Daura Jorge, F. G., Cantor, M., Ingram, S. N., Lusseau, D., & Simões Lopes, P. C. (2012). The structure of a bottlenose dolphin society is coupled to a unique foraging cooperation with artisanal fishermen. Biology Letters, 8(5), 702–705. https://doi.org/10.1098/rsbl.2012.0174

7 Cantor, M., Farine, D. R., & Daura‑Jorge, F. G. (2023). Foraging synchrony drives resilience in human–dolphin mutualism. Proceedings of the National Academy of Sciences, 120(6), e2207739120. https://doi.org/10.1073/pnas.2207739120

8 Jensen, A. M. (2012). The material culture of Iñupiat whaling: An ethnographic and ethnohistorical perspective. Arctic Anthropology, 49(2), 143–161. https://doi.org/10.1353/arc.2012.0020

9 Description of the USA Aboriginal Subsistence Hunt: Alaska. (n.d.). Iwc.int. https://iwc.int/management-and-conservation/whaling/aboriginal/usa/alaska

10 Derville, S., Buell, T. V., Corbett, K. C., Hayslip, C., & Torres, L. G. (2023). Exposure of whales to entanglement risk in Dungeness crab fishing gear in Oregon, USA. Biological Conservation, 281, 109989. https://doi.org/10.1016/j.biocon.2023.109989

11 Silber, G. K., Weller, D. W., Reeves, R. R., Adams, J. D., & Moore, T. J. (2021). Co‑occurrence of gray whales and vessel traffic in the North Pacific Ocean. Endangered Species Research, 44, 177–201. https://doi.org/10.3354/esr01093

12 Sullivan, F. A., & Torres, L. G. (2018). Assessment of vessel disturbance to gray whales to inform sustainable ecotourism. Journal of Wildlife Management, 82(5), 896–905. https://doi.org/10.1002/jwmg.21462

13 Sprogis, K. R., Videsen, S., & Madsen, P. T. (2020). Vessel noise levels drive behavioural responses of humpback whales with implications for whale‑watching. eLife, 9, e56760. https://doi.org/10.7554/eLife.56760

14 Lagerquist, B. A., Palacios, D. M., Winsor, M. H., Irvine, L. M., Follett, T. M., & Mate, B. R. (2019). Feeding home ranges of Pacific Coast Feeding Group gray whales. Journal of Wildlife Management, 83(4), 925–937. https://doi.org/10.1002/jwmg.21642

15 GRANITE: Gray whale Response to Ambient Noise Informed by Technology and Ecology | Marine Mammal Institute | Oregon State University. (n.d.). Mmi.oregonstate.edu. https://mmi.oregonstate.edu/gemm-lab/granite-gray-whale-response-ambient-noise-informed-technology-ecology

16 Pirotta, E., Bierlich, K. C., New, L., Bird, C. N., Fernandez Ajó, A., Hildebrand, L., Buck, C. L., Hunt, K. E., Calambokidis, J., & Torres, L. G. (2025). Body size, nutritional state and endocrine state are associated with calving probability in a long‑lived marine species. Journal of Animal Ecology. Advance online publication. https://doi.org/10.1111/1365-2656.70068

17 Bierlich, K. C., Kane, A., Hildebrand, L., Bird, C. N., Fernandez Ajó, A., Stewart, J. D., Hewitt, J., Hildebrand, I., Sumich, J., & Torres, L. G. (2023). Downsized: Gray whales using an alternative foraging ground have smaller morphology. Biology Letters, 19(7), 20230043. https://doi.org/10.1098/rsbl.2023.0043

New GEMM Lab study indicates troubled times for PCFG gray whales

Dr. Enrico Pirotta (CREEM, University of St Andrews) and Dr. Leigh Torres (GEMM Lab, MMI, OSU)

The health of animals affects their ability to survive and reproduce, which, in turn, drives the dynamics of populations, including whether their abundance trends up or down. Thus, understanding the links between health and reproduction can help us evaluate the impact of human activities and climate change on wildlife, and effectively guide our management and conservation efforts. In long-lived species, such as whales, once a decline in population abundance is detected, it can be too late to reverse the trend, so early warning signals are needed to indicate how these populations are faring.

We worked on this complex issue in a study that was recently published in the Journal of Animal Ecology. In this paper, we developed a new statistical approach to link three key components of the health of a Pacific Coast Feeding Group (PCFG) gray whale (namely, its body size, body condition, and stress levels) to a female’s ability to give birth to a calf. We were able to inform these metrics of whale health using an eight-year dataset derived from the GRANITE project of aerial images from drones for measurements of body size and condition, and fecal samples for glucocorticoid hormone analysis as an indicator of stress. We combined these data with observations of females with or without calves throughout the PCFG range over our study period.

We found that for a female to successfully have a calf, she needs to be both large and fat, as these factors indicate if the female has enough energy stored to support reproduction that year (Fig. 1). Remarkably, we also found indication that females with particularly high stress hormone levels may not get pregnant in the first place, which is the first demonstration of a link between stress physiology and vital rates in a baleen whale, to our knowledge.

Figure 1. Taken from Pirotta et al. (2025), Fig. 5. Combined relationship of PCFG gray whale length and nutritional state (combination of body size and condition) in the previous year with calving probability, colored by whether the model estimated an individual to have calved or not at a given reproductive opportunity.

Our study’s findings are concerning given our previous research indicating that gray whales in this PCFG sub-group have been growing to shorter lengths over the last couple of decades (Pirotta et al. 2023), are thinner than animals in the broader Eastern North Pacific gray whale population (Torres et al, 2022), and show an increase in stress-related hormones when exposed to human activities (Lemos et al, 2022; Pirotta et al. 2023). Furthermore, in our recent study we also documented that there are fewer young individuals than expected for a growing or stable population (Fig. 2), which can be an indicator of a population in decline since there may not be many individuals entering the reproductive adult age groups. Altogether, our results act as early warning signals that the PCFG may be facing a possible population decline currently or in the near future.

Figure 2. Taken from Pirotta et al. (2025), Fig. 1. Age structure diagram for 139 PCFG gray whales in our dataset. Each bar represents the number of individuals of a given age in 2023, with the color indicating the proportion of individuals of that age for which age is known (vs. estimated from a minimum estimate following Pirotta, Bierlich, et al., 2024). The red line reports a smooth kernel density estimate of the distribution.

These findings are sobering news for Oregon residents and tourists who enjoy watching these whales along our coast every summer and fall. We have gotten to know many of these whales so well – like Scarlett, Equal, Clouds, Lunita, and Pacman, who you can meet on our IndividuWhale website – that we wonder how they will adapt and survive as their once reliable habitat and prey-base changes. We hope our work sparks collective and multifaceted efforts to reduce impacts on these unique PCFG whales, and that we can continue the GRANITE project for many more years to come to monitor these whales and learn from their response to change.

This work exemplifies the incredible value of long-term studies, interdisciplinary methods, and effective collaboration. Through many years of research on this gray whale group, we have collected detailed data on diverse aspects of their behavior, ecology and life history that are critical to understanding their response to disturbance and environmental change, which are both escalating in the study region. We are incredibly grateful to the following members of the PCFG Consortium for contributing sightings and calf observation data that supported this study: Jeff Jacobsen, Carrie Newell, NOAA Fisheries (Peter Mahoney and Jeff Harris), Cascadia Research Collective (Alie Perez), Department of Fisheries and Oceans, Canada (Thomas Doniol-Valcroze and Erin Foster), Mark Sawyer and Ashley Hoyland, Wendy Szaniszlo, Brian Gisborne, Era Horton.

Did you enjoy this blog? Want to learn more about marine life, research, and conservation? Subscribe to our blog and get a weekly alert when we make a new post! Just add your name into the subscribe box below!

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References:

Lemos, Leila S., Joseph H. Haxel, Amy Olsen, Jonathan D. Burnett, Angela Smith, Todd E. Chandler, Sharon L. Nieukirk, Shawn E. Larson, Kathleen E. Hunt, and Leigh G. Torres. “Effects of Vessel Traffic and Ocean Noise on Gray Whale Stress Hormones.” Scientific Reports 12, no. 1 (2022): 18580. https://dx.doi.org/10.1038/s41598-022-14510-5.

Pirotta, Enrico, K. C. Bierlich, Leslie New, Lisa Hildebrand, Clara N. Bird, Alejandro Fernandez Ajó, and Leigh G. Torres. “Modeling Individual Growth Reveals Decreasing Gray Whale Body Length and Correlations with Ocean Climate Indices at Multiple Scales.” Global Change Biology 30, no. 6 (2024): e17366. https://doi.org/https://doi.org/10.1111/gcb.17366. https://onlinelibrary.wiley.com/doi/abs/10.1111/gcb.17366.

Pirotta, Enrico, Alejandro Fernandez Ajó, K. C. Bierlich, Clara N Bird, C Loren Buck, Samara M Haver, Joseph H Haxel, Lisa Hildebrand, Kathleen E Hunt, Leila S Lemos, Leslie New, and Leigh G Torres. “Assessing Variation in Faecal Glucocorticoid Concentrations in Gray Whales Exposed to Anthropogenic Stressors.” Conservation Physiology 11, no. 1 (2023). https://dx.doi.org/10.1093/conphys/coad082.

Torres, Leigh G., Clara N. Bird, Fabian Rodríguez-González, Fredrik Christiansen, Lars Bejder, Leila Lemos, Jorge Urban R, et al. “Range-Wide Comparison of Gray Whale Body Condition Reveals Contrasting Sub-Population Health Characteristics and Vulnerability to Environmental Change.” Frontiers in Marine Science 9 (2022). https://doi.org/10.3389/fmars.2022.867258. https://www.frontiersin.org/article/10.3389/fmars.2022.867258