Good enough to eat: Dynamics of krill prey quality

By Rachel Kaplan, PhD candidate, Oregon State University College of Earth, Ocean, and Atmospheric Sciences and Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

The Northern California Current region feeds a taxonomically diverse suite of top predators, including numerous species of seabirds, fish, and marine mammals. Baleen whales such as blue, fin, and humpback whales make this productive area a long stop on their seasonal migration, drawn in large part by abundant krill, a shrimp-like zooplankton that serves as an important prey item.

Aspects of both quality and quantity determine whether a prey resource is advantageous for a predator. In the case of whales, sheer biomass is key. It takes a lot of tiny krill to sustain a large whale – literally tons for a blue whale’s daily diet (Goldbogen et al., 2015). Baleen whales are such big eaters that they actually reshape the ocean ecosystem around them (Savoca et al., 2021).

Figure 1. A blue whale lunge feeds on a shallow krill swarm. Read more here.

But the quality of prey, in addition to its quantity, is crucial to ener­getic profitability, and baleen whales must weigh both elements in their foraging decisions. The outcome of those calculations manifest in the diverse feeding strategies that whales employ across ecosystems. In the California Current region, blue whales prefer­entially target the larger, more lipid-rich krill Thysanoessa spinifera (Fiedler et al., 1998). In Antarctica, humpback whales target larger and reproductive krill with higher energetic value, if these extra-juicy varieties are available (Cade et al., 2022). Prey-switching, a strategy in which animals target prey based on relative availability, allows fin whales to  have a more broad diet than blue whales, which are obligate krill predators.

So, what makes krill of high enough quality for a whale to pursue – or low enough quality to ignore? Krill are widely distributed across the NCC region, so why do foraging whales target one krill patch over another?

That whale of a question combines behavior, foraging theory, biochemistry, physics, climate, and more. One key aspect is the composition of a given prey item. Just as for human diet, nutrients, proteins, and calories are where the rubber hits the road in an animal’s energetic budget. The energy density of prey items sets the cost of living for cetaceans, and shapes the foraging strategies they use (Goldbogen et al., 2015; Spitz et al., 2012). In the NCC, T. spinifera krill are more lipid-rich than Euphausia pacifica (Fisher et al., 2020). Pursuing more energy dense prey increases the profitability of a given mouthful and helps a whale offset the energy expended to earn it, including the costly hunt for prey on the foraging grounds (Videsen et al., 2023).

Krill are amazingly dynamic animals in their own right, and they have evolved life history strategies to accommodate a broad range of ocean conditions. They can even exhibit “negative growth,” shrinking their body length in response to challenging conditions or poor food quality. This plasticity in body size can allow krill to survive lean times – but from the perspective of a hungry whale, this strategy also shrinks the available biomass into smaller packages (Robertson & Bjorkstedt, 2020).

One reason why krill are such advantageous prey type for baleen whales is their tendency to aggregate into dense swarms that may contain hundreds of thousands of individuals. The large body size of baleen whales requires them to feed on such profitable patches (Benoit-Bird, 2024). The packing density of krill within aggregations determines how many a whale can capture in one mouthful, and drives patch selection, such as for blue whales in Antarctica (Miller 2019).

Figure 2. The dense swarms formed by krill make them a prime target for many predators, including these juvenile Pacific sardines. (Photo: Richard Herrmann)

However, even the juiciest, densest krill won’t benefit a foraging whale if the energy required to consume it outweighs the gains. The depth of krill in the water column shapes the acrobatic foraging maneuvers blue whales use to feed (Goldbogen et al., 2015), and is a key driver of patch selection (Miller et al., 2019). The horizontal distance between the whale and a new krill patch is important too. Foraging humpback whales adapt their movements to the hierarchical structure of the preyfield, and feeding on neighboring prey schools can reduce the energy and time expended during interpatch travel, increasing net foraging gain (Kirchner et al., 2018).

Prey quality is dynamic, shaped by environmental conditions, extreme events, and climate change processes (Gomes et al., 2024). We can’t yet fully predict how change will alter prey and predator relationships in the NCC region (Muhling et al., 2020), making every step toward understanding prey dynamics relative to environmental variability key to anticipating how whales will fare in an unknown future (Hildebrand et al., 2021). If you are what you eat, then learning more about krill prey quality will give us unique insights into the baleen whales that come from far and wide to the NCC foraging grounds.



Benoit-Bird, K. J. (2024). Resource Patchiness as a Resolution to the Food Paradox in the Sea. The American Naturalist, 203(1), 1–13.

Cade, D. E., Kahane-Rapport, S. R., Wallis, B., Goldbogen, J. A., & Friedlaender, A. S. (2022). Evidence for Size-Selective Predation by Antarctic Humpback Whales. Frontiers in Marine Science, 9, 747788.

Fiedler, P. C., Reilly, S. B., Hewitt, R. P., Demer, D., Philbrick, V. A., Smith, S., Armstrong, W., Croll, D. A., Tershy, B. R., & Mate, B. R. (1998). Blue whale habitat and prey in the California Channel Islands. Deep Sea Research Part II: Topical Studies in Oceanography, 45(8–9), 1781–1801.

Fisher, J. L., Menkel, J., Copeman, L., Shaw, C. T., Feinberg, L. R., & Peterson, W. T. (2020). Comparison of condition metrics and lipid content between Euphausia pacifica and Thysanoessa spinifera in the northern California Current, USA. Progress in Oceanography, 188.

Goldbogen, J. A., Hazen, E. L., Friedlaender, A. S., Calambokidis, J., DeRuiter, S. L., Stimpert, A. K., & Southall, B. L. (2015). Prey density and distribution drive the three‐dimensional foraging strategies of the largest filter feeder. Functional Ecology, 29(7), 951–961.

Gomes, D. G. E., Ruzicka, J. J., Crozier, L. G., Huff, D. D., Brodeur, R. D., & Stewart, J. D. (2024). Marine heatwaves disrupt ecosystem structure and function via altered food webs and energy flux. Nature Communications, 15(1), 1988.

Hildebrand, L., Bernard, K. S., & Torres, L. G. (2021). Do Gray Whales Count Calories? Comparing Energetic Values of Gray Whale Prey Across Two Different Feeding Grounds in the Eastern North Pacific. Frontiers in Marine Science, 8.

Kirchner, T., Wiley, D., Hazen, E., Parks, S., Torres, L., & Friedlaender, A. (2018). Hierarchical foraging movement of humpback whales relative to the structure of their prey. Marine Ecology Progress Series, 607, 237–250.

Miller, E. J., Potts, J. M., Cox, M. J., Miller, B. S., Calderan, S., Leaper, R., Olson, P. A., O’Driscoll, R. L., & Double, M. C. (2019). The characteristics of krill swarms in relation to aggregating Antarctic blue whales. Scientific Reports, 9(1), 16487.

Muhling, B. A., Brodie, S., Smith, J. A., Tommasi, D., Gaitan, C. F., Hazen, E. L., Jacox, M. G., Auth, T. D., & Brodeur, R. D. (2020). Predictability of Species Distributions Deteriorates Under Novel Environmental Conditions in the California Current System. Frontiers in Marine Science, 7.

Robertson, R. R., & Bjorkstedt, E. P. (2020). Climate-driven variability in Euphausia pacifica size distributions off northern California. Progress in Oceanography, 188.

Savoca, M. S., Czapanskiy, M. F., Kahane-Rapport, S. R., Gough, W. T., Fahlbusch, J. A., Bierlich, K. C., Segre, P. S., Di Clemente, J., Penry, G. S., Wiley, D. N., Calambokidis, J., Nowacek, D. P., Johnston, D. W., Pyenson, N. D., Friedlaender, A. S., Hazen, E. L., & Goldbogen, J. A. (2021). Baleen whale prey consumption based on high-resolution foraging measurements. Nature, 599(7883), 85–90.

Spitz, J., Trites, A. W., Becquet, V., Brind’Amour, A., Cherel, Y., Galois, R., & Ridoux, V. (2012). Cost of Living Dictates what Whales, Dolphins and Porpoises Eat: The Importance of Prey Quality on Predator Foraging Strategies. PLoS ONE, 7(11), e50096.

Videsen, S. K. A., Simon, M., Christiansen, F., Friedlaender, A., Goldbogen, J., Malte, H., Segre, P., Wang, T., Johnson, M., & Madsen, P. T. (2023). Cheap gulp foraging of a giga-predator enables efficient exploitation of sparse prey. Science Advances, 9(25), eade3889.

First Flight

By Lindsay Wickman, Postdoc, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

I’ve had the privilege of attending several marine mammal surveys aboard ships at sea, but I had never surveyed for marine mammals from the air. So, when given the opportunity to participate in ongoing aerial surveys off the Oregon Coast with the US Coastguard’s helicopter fleet, I enthusiastically said yes. As Craig Hayslip, a Faculty Research Assistant with the Marine Mammal Institute, prepared me for my first helicopter survey, I was all excitement and no nerves. That is, until he explained the seating arrangement.

“There are two types of helicopters you’ll be flying on, and because of the seating arrangement in the Jayhawk, we fly with the door open when surveying for whales – it’s the only way to get a sufficient view,” Craig casually explained. I stared at the iPad I would use for recording data and imagined it flying through that open door and toward the sea, while I looked on flustered and helpless. Sensing my worry, Craig quickly showed me a set of straps that attached to the iPad, so it could be secured to one of my legs.

In addition to ensuring the iPad stayed in the aircraft, the straps also meant my hands would still be free to handle the camera (to aid in species identification), and a small tool called a geometer (developed by Pi Techology). By lining up the whale sighting in the sight of the geometer, the observer can record the angle between the aircraft and the sighting. Since we also know the height of the helicopter (we fly at a constant altitude of 500 feet), this angle can be used to calculate horizontal distance from the aircraft, allowing an accurate location to be estimated for each sighting.

My first flight was from Warrenton, Oregon, a four-hour drive north from the Hatfield Marine Science Center in Newport. Once at the airport, our first stop was to head to the flight operations office (a.k.a. “Ops”), who set us up with proper clothing and headgear for the flight. As we checked in, rock music played on a speaker while uniformed Coast Guardsmen serviced a helicopter in the hangar. I started to feel like a cool insider, until I clumsily donned the canvas flight suit and tried on several helmets. Suddenly several pounds heavier, all my movements became very awkward.

Lindsay outside the hangar wearing flight gear, in front of the survey’s helicopter. Photo by Craig Hayslip.

After my safety briefing, the entire crew gathered for a pre-flight meeting. We discussed weather conditions, did a wellness check, and discussed the flight’s mission. The conversation also included a brief overview of our scientific aims – why exactly were we looking for whales?

Craig briefly described the research project we were contributing to, titled Overlap Predictions About Large whales (OPAL). The main goal of this project is to better understand the overlap between whales and fisheries, with the aim of reducing entanglement risk. Fishing methods that use fixed, vertical lines in the water column, like the Dungeness crab fishery, can entangle whales as they migrate and feed along Oregon’s coastline. Since reports of whale entanglements have increased on the West Coast in the last 10 years, managing this threat is essential to ensure both the health of whale populations and the stability of Oregon’s crab fishery. Preventing these entanglements requires an understanding of where whales are distributed along the coast, as well as the times of year overlap with fisheries is most likely to occur. The OPAL project isn’t just mapping whale sightings, though. By using models to correlate whale sightings with oceanographic conditions, OPAL is also aiming to predict where whales are likely to occur.

After explaining the mission, the crew had to reach a consensus on both the level of “risk” in the mission and its level of “gain.” For a whale survey flight, risk was deemed low, with medium gain. While I initially felt mild offence that our scientific work was deemed to have just “medium” gain, I quickly reminded myself that when the crew is not flying scientists around, they are literally saving human lives. It was also a reminder that our whale surveys could easily be interrupted if necessary – Craig had mentioned several instances where flights were diverted to assist in rescue or medical emergencies.

With the briefing over, each of us had to consent to the flight plan by saying, “I accept this mission.” I’d heard this phrase from secret agents and soldiers in movies, but never from a marine scientist. I felt out of place saying them at first, but the words undeniably helped me establish a self-assured confidence I would give the survey my 100%.

Finally, it was time to head out of the hangar and to the aircraft. With both a pair of earplugs and my flight helmet on, the whirring of the blades was just a soft hum. I couldn’t hear speech, so we all relied on hand signals to communicate until our headsets were connected to the aircraft. The crew helped make sure I correctly put on my seatbelt harness, which had not just one, but five buckles. While I still felt some mild concern for the iPad strapped to my leg, at least I knew I wouldn’t fall out.

Lindsay holds up the geometer during the flight. Photo by Craig Hayslip.

Craig helped ensure I had all the equipment set up properly: the iPad’s survey program, the GPS tracking, and the computer recording the geometer’s measurements. Soon after, the helicopter slowly rose, hovering above the runway, before turning and heading towards the coast at speed. My stomach dropped slightly, my ears popped, and cold air rushed through the open door. I looked out at the Columbia River as it stretched toward the coastline and out to sea, and I couldn’t stop smiling.

A rainbow mid-air. Photo by Craig Hayslip.

As we approached the ocean, my attention shifted back to the mission, and I started scanning the surface for whale blows. With the large helmet on, I noticed the camera and geometer were much more difficult to use, so I also made “practice sightings” of passing boats and buoys. It didn’t take long before my first real whale sighting though – two gray whales (Eschrichtius robustus). Over the next two hours, I saw four more gray whales, and six more whales I was unable to identify due to distance. With each sighting, I had to act fast to make each geometer recording. The helicopter travels at a speed of 90 knots and whales can disappear soon after surfacing.

Two hours of flying with the door open meant my nose was running and my typing skills were worsening due to cold fingers. As exciting as it was to spot whales from the air, I was a little relieved when we arrived back at the airport and I could warm back up. Luckily, my nightmare of losing an iPad from the helicopter did not come true, and I was returning home with another survey to add to over 200 (and counting!) helicopter surveys completed for the OPAL project. Four different flights covering different parts of the Oregon coast are completed each month, so I know I have more flights to look forward to. After a successful first mission, I feel ready to take on my next flight.

The four flight routes completed monthly for the OPAL project. Helicopter flights are enabled through a partnership with the US Coastguard.

If you’d like to learn more about the OPAL research project, check out these past blog posts:

A Matter of Time: Adaptively Managing the Timescales of Ocean Change and Human Response

The pathway to advancing knowledge of rorqual whale distribution off Oregon

From land, sea,… and space: searching for whales in the vast ocean

The ups and downs of the ocean

Recent publications presenting findings from the first two years of OPAL include:

Derville, S., Barlow, D. R., Hayslip, C., & Torres, L. G. (2022). Seasonal, Annual, and Decadal Distribution of Three Rorqual Whale Species Relative to Dynamic Ocean Conditions Off Oregon, USA. Frontiers in Marine Science, 9.

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, reveals distinctive spatio-temporal and climatic patterns. Biological Conservation, 281.

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