How fat do baleen whales get? Recent publication shows how humpback whales increase their body condition over the foraging season. 

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

Traveling across oceans takes a lot of energy. Most baleen whales use stored energy acquired on their summer foraging grounds to support the costs of migration to and reproduction on their winter breeding grounds. Since little, if any, feeding takes place during the migration and winter season, it is essential that baleen whales obtain enough food to increase their fat reserves to support reproduction. As such, baleen whales are voracious feeders, and they typically depart the foraging grounds much fatter than when they had arrived. 

So, how fat do baleen whales typically get by the end of the foraging season, and how does this differ across reproductive classes, such as a juvenile female vs. a pregnant female? Understanding these questions is key for identifying what a typical “healthy” whale looks like, information which can then help scientists and managers monitor potential impacts from environmental and anthropogenic stressors. In this blog, I will discuss a recent publication in Frontiers in Marine Science ( that is from my PhD dissertation with the Duke University Marine Robotics and Remote Sensing (MaRRS) Lab, and also includes GEMM lab members Allison Dawn and Clara Bird. In this study, we analyzed how humpback whales (Megaptera novaeangliae) along the Western Antarctic Peninsula (WAP) increase their fat reserves throughout the austral summer foraging season (Bierlich et al., 2022). This work also helps provide insight to the GEMM Lab’s GRANITE project (Gray whale Response to Ambient Noise Informed by Technology and Ecology), where we are interested in how Pacific Coast Feeding Group (PCFG) gray whales increase their energy reserves in response to environmental variability and increasing human activities. 

Eastern South Pacific humpback whales, identified as Stock G by the International Whaling Commission, travel over 16,000 km between summer foraging grounds along the WAP and winter breeding grounds between Ecuador and Costa Rica (Fig. 1). Like most baleen whales, Stock G humpback whales were heavily exploited by 20th century commercial whaling. Recent evidence suggests that this population is recovering, with an estimated increase in population size of ~7,000 individuals in 2000 to ~19,107 in 2020 (Johannessen et al., 2022). 

However, there are long-term concerns for this population. The WAP is one of the fastest warming regions on the planet, and regional populations of krill, an important food source for humpback whales, have declined steeply over the past half-century. Additionally, the WAP has seen a rapid expansion of human activities, such as tourism and krill fishing. Specifically, the WAP has experienced an increase in tourism from a total of 6,700 visitors from 59 voyages in 1990 to 73,000 visitors from 408 voyages in 2020, which may be causing increased stress levels amongst Stock G (Pallin et al., 2022). Furthermore, the krill fishery has increased harvest activities in key foraging areas for humpback whales (Reisinger et al., 2022). Understanding how humpback whales increase their energy reserves over the course of the foraging season can help researchers establish a baseline to monitor future impacts from climate change and human activities. This work also provides an opportunity for comparisons to other baleen whale populations that are also exposed to multiple stressors, such as the PCFG gray whales off the Newport Coast who are constantly exposed to vessel traffic and at risk of entanglement from fishing gear. 

Figure 1. The migration route of the Stock G humpback whale population. Figure adapted from Whales of the Antarctic Peninsula Report, WWF 2018.

To understand how humpback whales increase their energy reserves throughout the foraging season, we collected drone imagery of whales along the WAP between November and June, 2017-2019 (Fig. 2). We used these images to measure the length and width of the whale to estimate body condition, which represents an animal’s relative energy reserve and can reflect foraging success (see previous blog). We collected drone imagery from a combination of research stations (Palmer Station), research vessels (Laurence M. Gould), and tour ships (One Ocean Expeditions). We used several different drones types and accounted for measurement uncertainty associated with the camera, focal length lens, altitude, and altimeter (barometer/LiDAR) from each drone (see previous blog and Bierlich et al., 2021a, 2021b). We also took biopsy samples to identify the sex of each individual and to determine if females were pregnant or not. 

Figure 2. Two humpbacks gracefully swimming in the chilly water along the Western Antarctic Peninsula. Photo taken by KC Bierlich & the Duke University Marine Robotics and Remote Sensing (MaRRS) Lab.

Our final dataset included body condition measurements for 228 total individuals. We found that body condition increased linearly between November and June for each reproductive class, which included calves, juvenile females, juvenile whales of unknown sex, lactating females, mature whales of unknown sex, and non-pregnant females (Fig. 3). This was an interesting finding because a recent publication analyzing tagged whales from the same population found that humpback whales have high foraging rates in early season that then significantly decrease by February and March (Nichols et al., 2022). So, despite these reduced foraging rates throughout the season, humpback whales continue to gain substantial mass into the late season. This continued increase in body condition implies a change in krill abundance and/or quality into the late season, which may compensate for the lower feeding rates. For example, krill density and biomass increases by over an order of magnitude across the season (Reiss et al., 2017) and their lipid content increases by ~4x (Hagen et al., 1996). Thus, humpback whales likely compensate for their lower feeding rates by feeding on denser and higher quality krill, ultimately increasing their efficiency in energy deposition. 

Figure 3. Body condition, here measured as Body Area Index (BAI), increases linearly for each reproductive class across the austral summer foraging season (Nov – June) for humpback whales along the Western Antarctic Peninsula. The shading represents the uncertainty around the estimated relationship. The colors represent the month of data collection.

We found that body condition increase varied amongst reproductive classes. For example, lactating females had the poorest measures of body condition across the season, reflecting the high energetic demands of nursing their calves (Fig. 3). Conversely, non-pregnant females had the highest body condition at the start of the season compared to all the other classes, likely reflecting the energy saved and recovered by skipping breeding that year.  Calves, juvenile whales, and mature whales all reached similar levels of body condition by the end of the season, though mature whales will likely invest most of their energy stores toward reproduction, whereas calves and juveniles likely invest toward growth. We also found a positive relationship between the total length of lactating females and their calves, suggesting that bigger moms have bigger calves (Fig. 4). A similar trend has also been observed in other baleen whale species including southern and North Atlantic right whales (Christiansen et al., 2018; Stewart et al., 2022).

Figure 4. Big mothers have big calves. Total length (TL) measurement between mother-calf pairs. The bars around each point represents the uncertainty (95% highest posterior density intervals). The colors represent the month of data collection. The blue line represents the best fit from a Deming regression, which incorporate measurement uncertainty in both the independent (mother’s TL) and dependent variable (calf’s TL).

The results from the humpback study provide insight for my current work exploring how PCFG gray whales increase their energy reserves in relation to environmental variability and increasing human activities. Over the past seven years, the GEMM Lab has been collecting drone images of PCFG gray whales off the coast of Oregon to measure their body condition (see this GRANITE Project blog). Many of the individuals we encounter are seen across years and throughout the foraging season, providing an opportunity to evaluate how an individual’s body condition is influenced by environmental variation, stress levels, maturity, and reproduction. For example, we had nine total body condition measurements of a female PCFG whale named “Sole”, who had a curvilinear increase in body condition throughout the summer foraging season – a rapid increase in early season that slowed as the season progressed (Fig. 5). This raises many questions for us: is this how most PCFG whales typically increase their body condition during the summer? Is this increase different for pregnant or lactating females? How is this increase impacted by environmental variability or anthropogenic stressors? Repeated measurements of individuals, in addition to Sole, in different reproductive classes across different years will help us determine what body condition is considered a healthy range for gray whales. This is particularly important for monitoring any potential health consequences from anthropogenic stressors, such as vessel noise and traffic (see recent blog by GEMM Lab alum Leila Lemos). We are currently analyzing body condition measurements between 2016 – 2022, so stay tuned for upcoming results!

Figure 6. Body condition, here measured as Body Area Index (BAI), increases curvilinearly for “Sole”, a mature female Pacific Coat Feeding Group gray whale, imaged nine times along the Oregon coast in 2021. The colors represent the month of data collection. 


Bierlich, K. C., Hewitt, J., Bird, C. N., Schick, R. S., Friedlaender, A., Torres, L. G., et al. (2021a). Comparing Uncertainty Associated With 1-, 2-, and 3D Aerial Photogrammetry-Based Body Condition Measurements of Baleen Whales. Front. Mar. Sci. 8, 1–16. doi:10.3389/fmars.2021.749943.

Bierlich, K. C., Hewitt, J., Schick, R. S., Pallin, L., Dale, J., Friedlaender, A. S., et al. (2022). Seasonal gain in body condition of foraging humpback whales along the Western Antarctic Peninsula. Front. Mar. Sci. 9, 1–16. doi:10.3389/fmars.2022.1036860.

Bierlich, K., Schick, R., Hewitt, J., Dale, J., Goldbogen, J., Friedlaender, A., et al. (2021b). Bayesian approach for predicting photogrammetric uncertainty in morphometric measurements derived from drones. Mar. Ecol. Prog. Ser. 673, 193–210. doi:10.3354/meps13814.

Christiansen, F., Vivier, F., Charlton, C., Ward, R., Amerson, A., Burnell, S., et al. (2018). Maternal body size and condition determine calf growth rates in southern right whales. Mar. Ecol. Prog. Ser. 592, 267–281.

Hagen, W., Van Vleet, E. S., and Kattner, G. (1996). Seasonal lipid storage as overwintering strategy of Antarctic krill. Mar. Ecol. Prog. Ser. 134, 85–89. doi:10.3354/meps134085.

Johannessen, J. E. D., Biuw, M., Lindstrøm, U., Ollus, V. M. S., Martín López, L. M., Gkikopoulou, K. C., et al. (2022). Intra-season variations in distribution and abundance of humpback whales in the West Antarctic Peninsula using cruise vessels as opportunistic platforms. Ecol. Evol. 12, 1–13. doi:10.1002/ece3.8571.

Nichols, R., Cade, D. E., Kahane-Rapport, S., Goldbogen, J., Simpert, A., Nowacek, D., et al. (2022). Intra-seasonal variation in feeding rates and diel foraging behavior in a seasonally fasting mammal, the humpback whale. Open Sci. 9, 211674.

Pallin, L. J., Botero-Acosta, N., Steel, D., Baker, C. S., Casey, C., Costa, D. P., et al. (2022). Variation in blubber cortisol levels in a recovering humpback whale population inhabiting a rapidly changing environment. Sci. Rep. 12, 1–13. doi:10.1038/s41598-022-24704-6.

Reisinger, R., Trathan, P. N., Johnson, C. M., Joyce, T. W., Durban, J. W., Pitman, R. L., et al. (2022). Spatiotemporal overlap of baleen whales and krill fisheries in the Antarctic Peninsula region. Front. Mar. Sci. doi:doi: 10.3389/fmars.2022.914726.

Reiss, C. S., Cossio, A., Santora, J. A., Dietrich, K. S., Murray, A., Greg Mitchell, B., et al. (2017). Overwinter habitat selection by Antarctic krill under varying sea-ice conditions: Implications for top predators and fishery management. Mar. Ecol. Prog. Ser. 568, 1–16. doi:10.3354/meps12099.

Stewart, J. D., Durban, J. W., Europe, H., Fearnbach, H., Hamilton, P. K., Knowlton, A. R., et al. (2022). Larger females have more calves : influence of maternal body length on fecundity in North Atlantic right whales. Mar. Ecol. Prog. Ser. 689, 179–189. doi:10.3354/meps14040.

Memoirs from above: drone observations of blue, humpback, Antarctic minke, and gray whales

By KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

With the GRANITE field season officially over, we are now processing all of the data we collected this summer. For me, I am starting to go through all the drone videos to take snapshots of each whale to measure their body condition. As I go through these videos, I am reflecting on the different experiences I am fortunate enough to have with flying different drones, in different environments, over different species of baleen whales: blue, humpback, Antarctic minke, and now gray whales. Each of these species have a different morphological design and body shape (Woodward et al., 2006), which leads to different behaviors that are noticeable from the drone. Drones create immense opportunity to learn how whales thrive in their natural environments [see previous blog for a quick history], and below are some of my memories from above. 

I first learned how drones could be used to study the morphology and behavior of large marine mammals during my master’s degree at Duke University, and was inspired by the early works of John Durban (Durban et al., 2015, 2016) Fredrick Christiansen (Christiansen et al., 2016) and Leigh Torres (Torres et al., 2018). I immediately recognized the value and utility of this technology as a new tool to better monitor the health of marine mammals. This revelation led me to pursue a PhD with the Duke University Marine Robotics and Remote Sensing (MaRRS) Lab led by Dr. Dave Johnston where I helped further develop tools and methods for collecting drone-based imagery on a range of species in different habitats. 

When flying drones over whales, there are a lot of moving parts; you’re on a boat that is moving, flying something that is moving, following something that is moving. These moving elements are a lot to think about, so I trained hard, so I did not have to think about each step and flying felt intuitive and natural. I did not grow up playing video games, so reaching this level of comfort with the controls took a lot of practice. I practiced for hours over the course of months before my first field excursion and received some excellent mentorship and training from Julian Dale, the lead engineer in the MaRRS Lab. Working with Julian and the many hours of training helped me establish a solid foundation in my piloting skills and feel confident working in various environments on different species. 

Blue whales offshore of Monterey, California. 

In 2017 and 2018 I was involved in collaborative project with the MaRRS Lab and Goldbogen Lab at Stanford University, where we tagged and flew drones over blue whales offshore of Monterey, California. We traveled about an hour offshore and reliably found groups of blue whales actively feeding. Working offshore typically brought a large swell, which can often make landing the drone back into your field partner’s hands tricky as everything is bobbing up and down with the oscillations of the swell. Fortunately, we worked from a larger research vessel (~56 ft) and quickly learned that landing the drone in the stern helped dampen the effects of bobbing up and down. The blue whales we encountered often dove to a depth of around 200 m for about 20-minute intervals, then come to the surface for only a few minutes. This short surface period provided only a brief window to locate the whale once it surfaced and quickly fly over it to collect the imagery needed before it repeated its dive cycle. We learned to be patient and get a sense of the animal’s dive cycle before launch in order to time our flights so the drone would be in the air a couple of minutes before the whale surfaced. 

Once over the whales, the streamlined body of the blue whales was noticeable, with their small, high aspect ratio flippers and fluke that make them so well adapted for fast swimming in the open ocean (Fig. 1) (Woodward et al., 2006). I also noticed that because these whales are so large (often 21 – 24 m), I often flew at higher altitudes to be able fit them within the field of view of the camera. It was also always shocking to see how small the tagging boat (~8 m) looked when next to Earth’s largest creatures. 

Figure 1. Two blue whales surface after a deep dive offshore of Monterey, Ca. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03)

Antarctic minke whales and humpback whales along the Western Antarctic PeninsulaA lot of the data included in my dissertation came from work along the Western Antarctic Peninsula (WAP), which had a huge range of weather conditions, from warm and sunny days to cold and snowy/foggy/rainy/windy/icy days. A big focus was often trying to keep my hands warm, as it was often easier to fly without gloves in order to better feel the controls. One of the coldest days I remember was late in the season in mid-June (almost winter!) in Wilhemina Bay where ice completely covered the bay in just a couple hours, pushing the whales out into the Gerlache Strait; I suspect this was the last ice-free day of the season. Surprisingly though, the WAP also brought some of the best conditions I have ever flown in. Humpback and Antarctic minke whales are often found deep within the bays along the peninsula, which provided protection from the wind. So, there were times where it would be blowing 40 mph in the Gerlache Strait, but calm and still in the bays, such as Andvord Bay, which allowed for some incredible conditions for flying. Working from small zodiacs (~7 m) allowed us more maneuverability for navigating around or through the ice deep in the bays (Fig. 2) 

Figure 2. Navigating through ice-flows along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Flying over Antarctic minke whale was always rewarding, as they are very sneaky and can quickly disappear under ice flows or in the deep, dark water. Flying over them often felt like a high-speed chase, as their small streamlined bodies makes them incredibly quick and maneuverable, doing barrel rolls, quick banked turns, and swimming under and around ice flows (Fig. 3). There would often be a group between 3-7 individuals and it felt like they were playing tag with each other – or perhaps with me!  

Figure 3. Two Antarctic minke whales swimming together along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Humpbacks displayed a wide range of behaviors along the WAP. Early in the season they continuously fed throughout the entire day, often bubble net feeding in groups typically of 2-5 animals (Fig. 4). For as large as they are, it was truly amazing to see how they use their pectoral fins to perform quick accelerations and high-speed maneuvering for tight synchronized turns to form bubble nets, which corral and trap their krill, their main food source (Fig. 4) (Woodward et al., 2006). Later in the season, humpbacks switched to more resting behavior in the day and mostly fed at night, taking advantage of the diel vertical migration of krill. This behavior meant we often found humpbacks snoozing at the surface after a short dive, as if they were in a food coma. They also seemed to be more curious and playful with each other and with us later in the season (Fig. 5).

We also encountered a lot of mom and calf pairs along the WAP. Moms were noticeably skinny compared to their plump calf in the beginning of the season due to the high energetic cost of lactation (Fig. 6). It is important for moms to regain this lost energy throughout the feeding season and begin to wean their calves. I often saw moms refusing to give milk to their nudging calf and instead led teaching lessons for feeding on their own.

Figure 4. Two humpback whales bubble-net feeding early in the feeding season (December) along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)
Figure 5. A curious humpback whale dives behind our Zodiac along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)
Figure 6. A mom and her calf rest at the surface along the Western Antarctic Peninsula. Note how the mom looks skinnier compared to her plump calf, as lactation is the most energetically costly phase of the reproductive cycle. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Gray whales off Newport, Oregon

All of these past experiences helped me quickly get up to speed and jump into action with the GRANITE field team when I officially joined the GEMM Lab this year in June. I had never flown a DJI Inspire quadcopter before (the drone used by the GEMM Lab), but with my foundation piloting different drones, some excellent guidance from Todd and Clara, and several hours of practice to get comfortable with the new setup, I was flying over my first gray whale by day three of the job. 

The Oregon coast brings all sorts of weather, and some days I strangely found myself wearing a similar number of layers as I did in Antarctica. Fog, wind, and swell could all change within the hour, so I learned to make the most of weather breaks when they came. I was most surprised by how noticeably different gray whales behave compared to the blue, Antarctic minke, and humpback whales I had grown familiar with watching from above. For one, it is absolutely incredible to see how these huge whales use their low-aspect ratio flippers and flukes (Woodward et al., 2006) to perform low-speed, highly dynamic maneuvers to swim in very shallow water (5-10 m) so close to shore (<1m sometimes!) and through kelp forest or surf zones close to the beach. They have amazing proprioception, or the body’s ability to sense its movement, action, and position, as gray whales often use their pectoral fins and fluke to stay in a head standing position (see Clara Bird’s blog) to feed in the bottom sediment layer, all while staying in the same position and resisting the surge of waves that could smash them against the rocks (Video 1) . It is also remarkable how the GEMM Lab knows each individual whale based on natural skin marks, and I started to get a better sense of each whale’s behavior, including where certain individuals typically like to feed, or what their dive cycle might be depending on their feeding behavior. 

Video 1. Two Pacific Coast Feeding Group (PCFG) gray whales “head-standing” in shallow waters off the coast of Newport, Oregon. NOAA/NMFS permit #21678

I feel very fortunate to be a part of the GRANITE field team and to contribute to data collection efforts. I look forward to the data analysis phase to see what we learn about how the morphology and behavior of these gray whales to help them thrive in their environment. 


Christiansen, F., Dujon, A. M., Sprogis, K. R., Arnould, J. P. Y., and Bejder, L. (2016).Noninvasive unmanned aerial vehicle provides estimates of the energetic cost of reproduction in humpback whales. Ecosphere 7, e01468–18.

Durban, J. W., Fearnbach, H., Barrett-Lennard, L. G., Perryman, W. L., & Leroi, D. J. (2015). Photogrammetry of killer whales using a small hexacopter launched at sea. Journal of Unmanned Vehicle Systems3(3), 131-135.

Durban, J. W., Moore, M. J., Chiang, G., Hickmott, L. S., Bocconcelli, A., Howes, G., et al.(2016). Photogrammetry of blue whales with an unmanned hexacopter. Mar. Mammal Sci. 32, 1510–1515.

Torres, L. G., Nieukirk, S. L., Lemos, L., & Chandler, T. E. (2018). Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science5, 319.

Woodward, B. L., Winn, J. P., and Fish, F. E. (2006). Morphological specializations of baleen whales associated with hydrodynamic performance and ecological niche. J. Morphol. 267, 1284–1294.

Wildlife of the Western Antarctic Peninsula

Erin Pickett, MS Student, Fisheries and Wildlife Department, OSU

This time last week, I was on a research vessel crossing the Drake Passage. The Drake extends from the tip of the Western Antarctic Peninsula to South America’s Cape Horn, and was part of the route I was taking home from Antarctica. Over the past three months I have been working on a long-term ecological research (LTER) project based out of Palmer Station, a U.S. based research facility located on Anvers Island.


While in Antarctica, I was working on the cetacean component of the Palmer LTER project, which I’ve described in previous blog posts. In lieu of writing more about what it is like to work and live on the Antarctic Peninsula, I thought I’d share some photos with you. Working on the water everyday while searching for whales provided me with many opportunities to photograph the local wildlife. I hope you’ll enjoy a few of my favorite shots.