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.

Toward an enhanced understanding of large whale ecology: a standardized protocol to quantify hormones in whale blubber

Dr. Alejandro A. Fernández Ajó, Postdoctoral Scholar, Marine Mammal Institute – OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab.

Whales are exposed to an increasing number of human-induced stressors—ranging from pollution and bycatch to the impacts of climate change on prey quality and distribution. Understanding how these factors affect whale health is critical for their conservation. The use of alternative approaches (i.e., alternative to blood samples) for gathering physiological information on large whales using a variety of non-lethal and non to minimally invasive sample matrices (i.e., blubber biopsies, blow, and fecal samples) provides a window into their endocrine state, allowing researchers to assess how these animals respond to both short-term and long-term stressors, and assess their reproductive and nutritional status. However, a lack of standardized protocols might hinder the comparability of results across studies, making it difficult to draw broad conclusions about the health and reproductive parameters of different whale populations.

Dr. Logan Pallin and I organized a lab exchange, funded by The Company of Biologists, to start a new collaboration aimed at bridging this gap by validating and standardizing methods for endocrine assessments in whale blubber. This is not just a technical exercise; it is a foundational step towards building equity and capacity in laboratories worldwide to conduct reliable and comparable endocrine assessments, enhancing the opportunities for multi-lab collaborations. Through this exchange, we aim to consolidate a standardized approach that will yield consistent results between laboratories, enabling better comparisons across different large whale populations. Hosted by the University of California Santa Cruz Biotelemetry and Behavioral Ecology Lab (UCSC-BTBEL Lab) under the mentorship of Dr. Logan Pallin, this experience is instrumental in advancing my research on large whale ecology and conservation.

Dr. Logan Pallin and Alejandro Fernandez Ajó conducting hormone extractions from gray whale blubber samples (left). Preparing a microtiter assay plate for hormone quantification in blubber (right).

During this exchange at the BBE Lab, I had the privilege of working closely with Dr. Logan Pallin, whose expertise in large whale endocrinology (particularly analyzing blubber biopsies) has been instrumental in shaping modern approaches to whale research. The lab’s cutting-edge equipment and Logan’s extensive experience with hormone extraction and quantification methods provided an ideal setting for refining our protocols. Our work focused on the extraction and quantification of progesterone from gray whale blubber samples provided by the Oregon State University Marine Mammal Stranding Network, part of MMI. These large blubber sections allow for repeated sub-sampling to ensure that the selected immunoassays reliably detect and measure the hormones of interest, while also assessing potential sources of variability when applying a standardized protocol. We initially focused our tests and validations on progesterone, as it is the precursor of all major steroid hormones and serves as an indicator of reproductive state in females.

A fieldwork day off Monterrey Bay, California with Dr. Logan Pallin, and PhD candidate Haley Robb. Blubber. Blubber biopsies can be obtained from free swimming whales with minimally invasive methods. From each sample we can derive multiple information about the reproductive status, genetics and overall health of the individuals.

The broader impact of our work
The successful validation and standardization of these protocols represents a significant advancement in whale conservation physiology. Once these methods are established, we plan to acquire funds to apply them to a larger collection of blubber samples. We hope to expand our work to include other species and regions, building a broader network of researchers dedicated to studying large whales in a rapidly changing world, and to assess hormone profiles in relation to factors like reproductive success, body condition, and exposure to stressors such as vessel traffic and environmental changes.

During our fieldwork in Monterey Bay, we had fascinating encounters with Minke whales (Balaenoptera acutorostrata, top left), a large group of Risso’s dolphins (Grampus griseus, bottom left), playful Humpbacks (Megaptera novaeangliae, top right), and a Blue whale (Balaenoptera musculus, no photo).

As I conclude this lab exchange, I am filled with excitement for the future. The knowledge and skills gained during this experience will undoubtedly shape the next phase of my research, allowing me to contribute more effectively to the conservation of these incredible animals. I look forward to applying these standardized methods to ongoing and future projects, and to continuing this fruitful collaboration with the BBE Lab. This journey has reinforced the importance of collaboration, standardization, and innovation in the field of conservation physiology. By working together, we can better understand the complex lives of large whales and take meaningful steps towards their protection in an increasingly challenging environment.

Acknowledgments: This exchange was made possible by the support of The Company of Biologists Traveling Fellowship Grant. I would like to thank Dr. Ari Friedlaender (BBE Lab PI) for facilitating this exchange, and Dr. Leigh Torres (GEMM Lab PI) and Dr. Lisa Balance (MMI director) for their support in helping me expand my collaboration network and skillsets. Special thanks to PhD student Haley Robb for her assistance in the laboratory and fieldwork, and a heartfelt thank you to Dr. Logan Pallin for generously sharing his knowledge and time.

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