Reflecting on a solitary journey surrounded by an incredible team

Clara Bird, PhD Candidate, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Graduate school is an odd phase of life, at least in my experience. You spend years hyperfocused on a project, learning countless new skills – and the journey is completely unique to you. Unlike high school or undergrad, you are on your own timeline. While you may have peers on similar timelines, at the end of day your major deadlines and milestone dates are your own. This has struck me throughout my time in grad school, and I’ve been thinking about it a lot lately as I approach my biggest, and final milestone – defending my PhD! 

I defend in just about two months, and to be honest, it’s very odd approaching a milestone like this alone. In high school and college, you count down to the end together. The feelings of anticipation, stress, excitement, and anticipatory grief that can accompany the lead-up to graduation are typically shared. This time, as I’m in an intense final push to the end while processing these emotions, most of the people around me are on their own unique timeline. At times grad school can feel quite lonely, but this journey would have been impossible without an incredible community of people.

A central contradiction of being a grad student is that your research is your own, but you need a variety of communities to successfully complete it. Your community of formal advisors, including your advisor and committee members, guide you along the way and provide feedback. Professors help you fill specific knowledge and skill gaps, while lab mates provide invaluable peer mentorship. Finally, fellow grad students share the experience and can celebrate and commiserate with you. I’ve also had the incredible fortune of having the community of the GRANITE team, and I’ve recently been reflecting on how special the experience has been.

To briefly recap, GRANITE stands for Gray whale Response to Ambient Noise Informed by Technology and Ecology (read this blog to learn more). This project is one of the GEMM lab’s long-running gray whale projects focused on studying gray whale behavior, physiology, and health to understand how whales respond to ocean noise. Given the many questions under this project, it takes a team of researchers to accomplish our goals. I have learned so much from being on the team. While we spend most of the year working on our own components, we have annual meetings that are always a highlight of the year. Our team is made up of ecologists, physiologists, and statisticians with backgrounds across a range of taxa and methodologies. These meetings are an incredible time to watch, and participate in, scientific collaboration in action. I have learned so much from watching experts critically think about questions and draw inspiration from their knowledge bases. It’s been a multi-year masterclass and a critically important piece of my PhD. 

The GRANITE team during our first in person meeting

These annual meetings have also served as markers of the passage of time. It’s been fascinating to observe how our discussions, questions, and ideas have evolved as the project progressed. In the early years, our presentations shared proposed research and our conversations focused on working out how on earth we were going to tackle the big questions we were posing. In parallel, it was so helpful to work out how I was going to accomplish my proposed PhD questions as part of this larger group effort. During the middle years, it was fun to hear progress updates and to learn from watching others go through their process too. In grad school, it’s easy to feel like your setbacks and stumbles are failures that reflect your own incompetence, but working alongside and learning from these scientists has helped remind me that setbacks and stumbles are just part of the process. Now, in the final phase, as results abound, it feels extra exciting to celebrate with this team that has watched the work, and me grow, from the beginning. 

The GRANITE team taking a beach walk after our second in person meeting.

We just wrapped up our last team meeting of the GRANITE project, and this year provided a learning experience in a phase of science that isn’t often emphasized in grad school. For graduate students, our work tends to end when we graduate. While we certainly think about follow-up questions to our studies, we rarely get the opportunity to follow through. In our final exams, we are often asked to think of next steps outside the constraints of funding or practicality, as a critical thinking exercise. But it’s a different skillset to dream up follow-up questions, and to then assess which of those questions are feasible and could come together to form a proposal. This last meeting felt like a cool full-story moment. From our earliest meetings determining how to answer our new questions, to now deciding what the next new questions are, I have learned countless lessons from watching this team operate. 

The GRANITE team after our third in person meeting.

There are a few overarching lessons I’ll take with me. First and foremost, the value of patience and kindness. As a young scientist stumbling up the learning curve of many skills all at once, I am so grateful for the patience and kindness I’ve been shown. Second, to keep an open mind and to draw inspiration from anything and everything. Studying whales is hard, and we often need to take ideas from studies on other animals. Which brings me to my third takeaway, to collaborate with scientists from a wide range of backgrounds who can combine their knowledges bases with yours, to generate better research questions and approaches to answering them.

I am so grateful to have worked with this team during my final sprint to the finish. Despite the pressure of the end nearing, I’m enjoying moments to reflect and be grateful. I am grateful for my teachers and peers and friends. And I can’t wait to share this project with everyone.

P.S. Interested in tuning into my defense seminar? Keep an eye on the GEMM lab Instagram (@gemm_lab) for the details and zoom link.

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New publication reveals gray whale habitat use patterns over three decades in the Northern California Current

By Dr. Dawn Barlow, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

The EMERALD project (Examining Marine mammal Ecology through Regional Assessment of Long-term Data) has reached a milestone with a recent publication detailing our findings on long-term gray whale distribution, abundance, and habitat use patterns (Barlow et al. 2024). The study is made possible by an incredible dataset. Every May-July since 1992, a team of observers surveyed the coastline between the Columbia River at the border between Oregon and Washington and San Francisco Bay, California for marbled murrelets, a seabird species of conservation concern. They drive a small vessel along pre-determined tracklines, and record observations of seabirds and marine mammals—not just marbled murrelets—and fortunately for us, that means there is a record of annual gray whale distribution and abundance patterns that spans over three decades.

The Crescent Coastal Research team collecting survey data. We are incredibly grateful to Craig Strong and the many folks who collected these valuable observations over the years!

We analyzed these valuable data using density surface modeling to better understand what drives gray whale distribution and abundance, what their habitat preferences are, and whether and how these occurrence patterns have changed over time. I am excited to share a few of our findings here!

Long-term, stable hotspots

The survey data revealed three main areas with consistently high gray whale density: the central Oregon Coast off Newport, Cape Blanco off Oregon’s south coast, and the mouth of the Klamath River in northern California. Despite fluctuations in how many whales were observed over the years, these areas have remained predicable hotspots for gray whales during their summer feeding season.

(A) Mean gray whale encounter rate (whales/kilometers surveyed) summarized by year, across all latitudes. (B) Mean gray whale encounter rate summarized by 1° latitude bin, across all years. White indicates times and locations with no survey effort. (C) Mean gray whale encounter rate summarized by year and 1° latitude bin. (D) Map of the study area, with region boundaries shown by the dashed lines, and major placenames denoted. Figure and caption reproduced from Barlow et al. 2024.

Key regional differences

Major features like prominent capes divide the California Current into different regions with distinct oceanographic characteristics. We found that gray whales showed different habitat preferences in the different regions. In the northern part of our study area between the Columbia River and Cape Blanco, we found that rocky bottom substrate was strongly related to areas of higher gray whale abundance, despite being far less available than soft, sandy bottom habitat. In the region between Cape Blanco and Cape Mendocino, gray whales were more abundant in areas south of prominent capes and in closer proximity to river estuaries.

Coastal upwelling and relaxation are key

Coastal upwelling—the process by which winds in the spring and summer push surface water offshore that is then replaced by cold, nutrient-rich water that is brought into the sunlight and drives an abundance of marine life—is a critically important influence in the oceanography, ecology, and biodiversity of our study region. But relaxation of those upwelling winds is also important for coastal species, as relaxation events allow the upwelled nutrients to be retained in the nearshore waters and enhance and aggregate local productivity and prey. We found that gray whale abundance was highest when there was a combination of both upwelling and relaxation events—a critical balance of “enough but not too much”—that seems to be optimal for gray whale feeding opportunities in nearshore waters.

You are what, where, and how you eat

Gray whales are incredibly flexible predators and have a wide range of prey items they are known to feed on. We found that throughout our study range, gray whales have different habitat preferences. As they spend their summers here to feed, these habitat preferences are linked to their foraging preferences. Off the central Oregon Coast, gray whales are known to feed on zooplankton that aggregate around rocky reefs and kelp forests (Hildebrand et al. 2022, 2024).

A gray whale surfaces in a patch of kelp, foraging around a rocky reef. UAS image credit: GEMM Lab.

Further south, in the region between Cape Blanco and Cape Mendocino that encompassed the long-term hotspot of gray whale sightings off the Klamath River, our models revealed different habitat preferences. In the soft-bottom habitat off the Klamath River, gray whales are known to do more benthic feeding, whereby they scoop up the seafloor and filter out the invertebrates in the sediment such as amphipods and cumaceans (Mallonée 1991, Jenkinson 2001).

A gray whale surfaces with a mouth full of muddy sediment, filtering out the invertebrate prey. UAS image credit: GEMM Lab.

These differences in regional habitat preferences and preferred prey likely relate to larger-scale phenomena as well. Indeed, when we looked at how gray whale abundance in different regions related to widespread warm or cool phases in the North Pacific Ocean, the responses differed by region. This aspect of the study indicates that what gray whales eat and where they forage influences how they respond to shifting environmental conditions and prey availability.

Conservation of an iconic nearshore predator

The unique mosaic of habitat characteristics throughout the Northern California Current summer feeding range of gray whales provides them the opportunity to gain the energetic stores they need to survive, reproduce, and migrate. Thus, the reliability of these resources has led them to return to these stable foraging hotspots year after year. Under climate change, one potential impact on upwelling systems is shifts in the intensity and location of upwelling (Bograd et al. 2023); in the Northern California Current, this could mean reduced relaxation events that we found are crucial for gray whales feeding in this habitat. Furthermore, these whales overlap with human activities such as vessel disturbance, entanglement and vessel strike risk, and ocean noise throughout the foraging season, and have to bear the consequences of these anthropogenic stressors (Sullivan & Torres 2018, Lemos et al. 2022, Pirotta et al. 2023) as they also navigate changing environmental conditions. Our study highlights the value of long-term monitoring to better understand present ecological patterns in the context of the past, which can be used to inform conservation management decisions for the future.

For more details, we invite you to read the full, open access publication here:

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Barlow DR, Strong CS, Torres LG (2024) Three decades of nearshore surveys reveal long-term patterns in gray whale habitat use, distribution, and abundance in the Northern California Current. Sci Rep 14:9352.

Bograd SJ, Jacox MG, Hazen EL, Lovecchio E, Montes I, Pozo Buil M, Shannon LJ, Sydeman WJ, Rykaczewski RR (2023) Climate Change Impacts on Eastern Boundary Upwelling Systems. Ann Rev Mar Sci 15:1–26.

Hildebrand L, Derville S, Hildebrand I, Torres LG (2024) Exploring indirect effects of a classic trophic cascade between urchins and kelp on zooplankton and whales. Sci Rep 14.

Hildebrand L, Sullivan FA, Orben RA, Derville S, Torres LG (2022) Trade-offs in prey quantity and quality in gray whale foraging. Mar Ecol Prog Ser 695:189–201.

Jenkinson RS (2001) Gray whale (Eschrichtius robustus) prey availability and feeding ecology in Northern California, 1999-2000. Humboldt State University

Lemos L, Haxel J, Olsen A, Burnett JD, Smith A, Chandler TE, Nieukirk SL, Larson SE, Hunt KE, Torres LG (2022) Effects of vessel traffic and ocean noise on gray whale stress hormones. Sci Rep 12:1–13.

Mallonée JS (1991) Behaviour of gray whales (Eschrichtius robustus) summering off the northern California coast, from Patrick’s Point to Crescent City. Can J Zool 69:681–690.

Pirotta E, Fernandez Ajó A, Bierlich KC, Bird CN, Buck CL, Haver SM, Haxel JH, Hildebrand L, Hunt KE, Lemos LS, New L, Torres LG (2023) Assessing variation in faecal glucocorticoid concentrations in gray whales exposed to anthropogenic stressors. Conserv Physiol 11:coad082.

Sullivan FA, Torres LG (2018) Assessment of vessel disturbance to gray whales to inform sustainable ecotourism. J Wildl Manage 82:896–905.

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.

Kelp to whales: New GEMM Lab publication explores indirect effects of a classic trophic cascade on gray whales

By Lisa Hildebrand, PhD candidate, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

As many of our avid readers already know, the Pacific Coast Feeding Group (PCFG) of gray whales employs a wide range of foraging tactics to feed on a number of different prey items in various benthic substrate types (Torres et al. 2018). One example foraging tactic is when PCFG whales, particularly when they are in the Oregon portion of their feeding range, forage on mysid shrimp in and near kelp beds on rocky reefs. We have countless drone video clips of whales weaving their large bodies through kelp and many photographs of whales coming to the surface to breath completely covered in kelp, looking more like a sea monster than a whale (Figure 1). So, when former intern Dylan Gregory made an astute observation during the 2018 TOPAZ/JASPER field season in Port Orford about a GoPro video the field team collected that showed many urchins voraciously feeding on an unhealthy-looking kelp stalk (Figure 2a), it made us wonder if and how changes to kelp forests may impact gray whales. 

Fig 1. Gray whale surfacing in a large kelp patch. Photograph captured under NOAA/NMFS research permit #16111. Source: GEMM Lab.

Kelp forests are widely used as a marine example of trophic cascades. Trophic cascades are trigged by the addition/removal of a top predator to/from a system, which causes changes further down the trophic chain. Sea urchins are common inhabitants of kelp forests and in a balanced, healthy system, urchin populations are regulated by predators as they behave cryptically by hiding in crevices in the reef and individual urchins feed passively on drift kelp that breaks off from larger plants. When we think about who controls urchins in kelp forests, we probably think of sea otters first. However, sea otters have been absent from Oregon waters for over a century (Kone et al. 2021), so who controls urchins here? The answer is the sunflower sea star (Figure 2b). Sunflower sea stars are large predators with a maximum arm span of up to 1 m! Unfortunately, a disease epidemic that started in 2013 known as sea star wasting disease caused 80-100% population decline of sunflower sea stars along the coastline between Mexico and Alaska (Harvell et al. 2019). Shortly thereafter, a record-breaking marine heatwave caused warm, nutrient-poor water conditions to persist in the northeast Pacific Ocean from 2014 to 2016 (Jacox et al. 2018). These co-occurring stressors caused unprecedented and long-lasting decline of a previously robust kelp forest in northern California (Rogers-Bennett & Catton 2019), where sea otters are also absent. Given the biogeographical similarity between southern Oregon and northern California and the observation made by Dylan in 2018, we decided to undertake an analysis of the eight years of data collected during the TOPAZ/JASPER project in Port Orford starting in 2016, to investigate the trends of four trophic levels (purple sea urchins, bull kelp, zooplankton, and gray whales) across space and time. The results of our study were published last week in Scientific Reports and I am excited to be able to share them with you today.

Every day during the TOPAZ/JASPER field season, two teams head out to collect data. One team is responsible for tracking gray whales from shore using a theodolite, while the other team heads out to sea on a tandem research kayak to collect prey data (Figure 3). The kayak team samples prey in multiple ways, including dropping a GoPro camera at each sampling station. When the project was first developed, the original goal of these GoPro videos was to measure the relative abundance of prey. Since the sampling stations occur on or near reefs that are shallow with dense surface kelp, traditional methods to assess prey density, such as using a boat with an echosounder, are not suitable options. Instead, GEMM Lab PI Leigh Torres, together with the first Master’s student on this project Florence Sullivan, developed a method to score still images extracted from the GoPro videos to estimate relative zooplankton abundance. However, after we saw those images of urchins feeding on kelp in 2018, we decided to develop another protocol that allowed us to use these GoPro videos to also characterize sea urchin coverage and kelp condition. Once we had occurrence values for all four species, we were able to dig into the spatiotemporal trends.

Figure 3. Map of Port Orford, USA study area showing the 10 kayak sampling stations (white circles) within the two study sites (Tichenor Cove and Mill Rocks). The white triangle represents the cliff top location where theodolite tracking of whales was conducted. Figure and caption taken from Hildebrand et al. 2024.

When we examined the trends for each of the four study species across years, we found that purple sea urchin coverage in both of our study sites within Port Orford increased dramatically within our study period (Figure 4). In 2016, the majority of our sampled stations contained no visible urchins. However, by 2020, we detected urchins at every sampling station. For kelp, we saw the reverse trend; in 2016 all sampling stations contained kelp that was healthy or mostly healthy. But by 2019, there were many stations that contained kelp in poor health or where kelp was absent entirely. Zooplankton and gray whales experienced similar temporal trends as the kelp, with their occurrence metrics (abundance and foraging time, respectively) having higher values at the start of our study period and declining steadily during the eight years. While the rise in urchin coverage across our study area occurred concurrently with the decrease in kelp condition, zooplankton abundance, and gray whale foraging, we wanted to explicitly test how these species are related to one another based on prior ecological knowledge.

Figure 4. Temporal trends of purple sea urchin coverage, bull kelp condition, relative zooplankton abundance, and gray whale foraging time by year across the eight-year study period (2016–2023), from the generalized additive models. The colored ribbons represent approximate 95% confidence intervals. Line types represent the two study sites, Mill Rocks (MR; solid) and Tichenor Cove (TC; dashed). All curves are statistically significant (P < 0.05). Figure and caption taken from Hildebrand et al. 2024.

To test whether urchin coverage had an effect on kelp condition, we hypothesized that increased urchin coverage would be correlated with reduced kelp condition based on the decades of research that has established a negative relationship between the two when a trophic cascade occurs in kelp forest systems. Next, we wanted to test whether kelp condition had an effect on zooplankton abundance and hypothesized that increased kelp condition would be correlated with increased zooplankton abundance. We based this hypothesis on several pieces of prior knowledge, particularly as they pertain to mysid shrimp: (1) high productivity within kelp beds provides food for mysids, including kelp zoospores (VanMeter & Edwards 2013), (2) current velocities are one third slower inside kelp beds compared to outside (Jackson & Winant 1983), which might support the retention of mysids within kelp beds since they are not strong swimmers, and (3) the kelp canopy may serve as potential protection for mysids from predators (Coyer 1984). Finally, we wanted to test whether both kelp condition and zooplankton abundance have an effect on gray whales and we hypothesized that increased values for both would be correlated with increased gray whale foraging time. While the reasoning behind our hypothesized correlation between zooplankton prey and gray whales is obvious (whales eat zooplankton), the reasoning behind the kelp-whale connection may not be. We speculated that since kelp habitat may aggregate or retain zooplankton prey, gray whales may use kelp as an environmental cue to find prey patches. 

When we tested our hypotheses through generalized additive models, we found that increased urchin coverage was significantly correlated with decreased kelp condition in both study sites, providing evidence that a shift from a kelp forest to an urchin barren may have occurred in the Port Orford area. Additionally, increased kelp condition was correlated with increased zooplankton abundance, supporting our hypothesis that kelp forests are an important habitat and resource for nearshore zooplankton prey. Interestingly, this relationship was bell-shaped in one of our two study sites, suggesting that there are other factors besides healthy bull kelp that influence zooplankton abundance, which likely include upwelling dynamics, habitat structure, and local oceanographic characteristics. For the whale model, we found that increased kelp condition was significantly correlated with increased gray whale foraging time, which may corroborate our hypothesis that gray whales use kelp as an environmental cue to locate prey. Zooplankton abundance was significantly correlated with gray whale foraging time in one of our two sites. Once again, this relationship was bell-shaped, which suggests other factors influence gray whale foraging time, including prey quality (Hildebrand et al. 2022) and density.

Figure 5. Effects derived from trophic path generalized additive models of purple sea urchin coverage on kelp condition (A), kelp condition on relative zooplankton abundance (B), and kelp condition and relative zooplankton abundance on gray whale foraging time (C). The colored ribbons represent approximate 95% confidence intervals. Line types represent the two study sites, Mill Rocks (MR; solid) and Tichenor Cove (TC; dashed). Curves with asterisks indicate statistically significant (P < 0.05) relationships. Figure and caption taken from Hildebrand et al. 2024.

Our results highlight the potential larger impacts of reduced gray whale foraging time as a result of these trophic dynamics may cause at the individual and population level. If an area that was once a reliable source of food (like Port Orford) is no longer favorable, then whales likely search for other areas in which to feed. However, if the areas affected by these dynamics are widespread, then individuals may spend more time searching for, and less time consuming, prey, which could have energetic consequences. While our study took place in a relatively small spatial area, the trophic dynamics we documented in our system may be representative of patterns across the PCFG range, given ecological and topographic similarities in habitat use patterns. In fact, in the years with the lowest kelp, zooplankton, and whale occurrence (2020 and 2021) in Port Orford, the GRANITE field team also noted low whale numbers and minimal surface kelp extent in the central Oregon field site off of Newport. However, ecosystems are resilient. We are hopeful that the dynamics we documented in Port Orford are just short-term changes and that the system will return to its former balanced state with less urchins, more healthy bull kelp, zooplankton, and lots of feeding gray whales.

If you are interested in getting a more detailed picture of our methods and analysis, you can read our open access paper here:

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Coyer, J. A. (1984). The invertebrate assemblage associated with the giant kelp, Macrocystis pyrifera, at Santa Catalina Island, California: a general description with emphasis on amphipods, copepods, mysids, and shrimps. Fishery Bulletin, 82(1), 55-66.

Harvell, C. D., Montecino-Latorre, D., Caldwell, J. M., Burt, J. M., Bosley, K., Keller, A., … & Gaydos, J. K. (2019). Disease epidemic and a marine heat wave are associated with the continental-scale collapse of a pivotal predator (Pycnopodia helianthoides). Science advances, 5(1), eaau7042.

Hildebrand, L., Sullivan, F. A., Orben, R. A., Derville, S., & Torres, L. G. (2022). Trade-offs in prey quantity and quality in gray whale foraging. Marine Ecology Progress Series, 695, 189-201.

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Significant others? Thinking beyond p-values in science

By Natalie Chazal, PhD student, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Scientific inquiry relies on quantifying how certain we are of the differences we see in observations. This means that we must look at phenomena based on probabilities that we calculate from observed data, or data that we collect from sampling efforts. Historically, p-values have served as a relatively ubiquitous tool for assessing the strength of evidence in support of a hypothesis. However, as our understanding of statistical methods evolves, so does the scrutiny surrounding the appropriateness and interpretation of p-values. In the realm of research, the debate surrounding the use of p-values for determining statistical significance has sparked some controversy and reflection within the academic community. 

What is a p-value?

To understand the debate itself, we need to understand what a p-value is. The p-value represents the probability of obtaining a result as extreme as, or more extreme than, the observed data, under the assumption that there is no true difference or relationship between groups or variables. Traditionally, a p-value below a predetermined threshold (often 0.05) is considered statistically significant, suggesting that the observed data are unlikely (i.e., a 5% probability) to have occurred by chance alone. Many statistical tests provide p-values, which gives us a unified framework for interpretation across a range of analyses.

To illustrate this, imagine a study aimed at investigating the effects of underwater noise pollution on the foraging behavior of gray whales. Researchers collect data on the diving behavior of gray whales in both noisy and quiet regions of the ocean.

Drawings of gray whales with tags (depicted by orange shapes) in quiet areas (left) and noisy areas (right). 

In this example, the researchers hypothesize that gray whales stop foraging and ultimately change their diving behavior in response to increased marine noise pollution. The data collected from this hypothetical scenario could come from tags equipped with sensors that record diving depth, duration, and location, allowing us to calculate the exact length of time spent foraging. Data would be collected from both noisy areas (maybe near shipping lanes or industrial sites) and quiet areas (more remote regions with minimal human activity). 

To assess the significance of the differences between the two noise regimes, researchers may use statistical tests like t-tests to compare two groups. In our example, researchers use a t-test to compare the average foraging time between whales in noisy and quiet regimes. The next step would be to define hypotheses about the differences we expect to see. The null hypothesis (HN) would be that there is no difference in the average foraging time (X) between noisy and quiet areas: 

Scenario where the noisy area does not elicit a behavioral response that can be detected by the data collected by the tags (orange shapes on whales back). The lower graph shows the distribution of the data (foraging time) for the noisy and the quiet areas. The means of this data (X) are not different. 

And the alternative hypothesis (HA) would be that there is a difference between the noisy and quiet areas: 

Scenario where the noisy area elicits a behavioral response (swimming more towards the surface instead of foraging) that can be detected by the data collected by the tags (orange shapes on whales back). The lower graph shows the distribution of the data (foraging time) for the noisy and the quiet areas. The means of this data (X) are different with the noisy mean foraging time (pink) being lower than the quiet mean foraging time (blue).

For now, we will skip over the nitty gritty of a t-test and just say that the researchers get a “t-score” that says whether or not there is a difference in the means (X) of the quiet and noisy areas. A larger t-score means that there is a difference in the means whereas a smaller t-score would indicate that the means are more similar. This t-score comes along with a p-value. Let’s say we get a t-score (green dot) that is associated with a p-value of 0.03 shown as the yellow area under the curve: 

The t-score is a test statistic that tells us how different the means of our observed data groups are from each other (green dot). The area under the t-distribution that is above the t-score is the p-value (yellow shaded area).

A p-value of 0.03 means that there is a 3% probability of obtaining these observed differences in foraging time between noisy and quiet areas purely by chance, which assumes that the null hypothesis is true (that there is no difference). We usually compare this p-value to a threshold value to say whether this finding is significant. We set this threshold before looking at the results of the test. If the threshold is above our value, like 0.05, then we can “reject the null hypothesis” and conclude that there is a significant difference in foraging time between noisy and quiet areas (green check mark scenario). On the flip-side, if the threshold that we set before our results is too low (0.01), then we will “fail to reject the null hypothesis” and conclude that there was no significant difference in foraging time between noisy and quiet areas (red check mark scenario). The reason that we don’t ever “accept the null” is because we are testing an alternative hypothesis with observations and if those observations are consistent with the null rather than the alternative, this is not evidence for the null because it could be consistent with a different alternative hypothesis that we are not yet testing for.

When our pre-set threshold to determine significance is above or greater than our p-value that was calculated we have enough evidence to ‘reject the null hypothesis’ (left figure) whereas if our p-value is lower or smaller than our calculated p-value, then we ‘fail to reject the null hypothesis’ (right figure).

In this example, the use of p-values helps the researchers quantify the strength of evidence for their hypothesis and determine whether the observed differences in gray whale behavior are likely to be meaningful or merely due to chance. 

The Debate

Despite its widespread use, the reliance on p-values has been met with criticism. Firstly, because p-values are so ubiquitous, it can be easy to calculate them with or without enough critical thinking or interpretation. This critical thinking should include an understanding of what is biologically relevant and avoid the trap of using binary language like significant or non-significant results instead of looking directly at the uncertainty of your results. One of the other most common misconceptions about p-values is that they can measure the direct probability of the null hypothesis being true. As amazing as that would be, in reality we can only use p-values to understand the probability of our observed data. Additionally, it’s common to conflate the significance or magnitude of the p-value with effect size (which is the strength of the relationship between the variables). You can have a small p-value for an effect that isn’t very large or meaningful, especially if you have a large sample size. Sample size is an important metric to report. Larger number of samples generally means more precise estimates, higher statistical power, increased generalizability, and higher possibility for replication.

Furthermore, in studies that require multiple comparisons (i.e. multiple statistical analyses are done in a single study), there is an increased likelihood of observing false positives because each test introduces a chance of obtaining a significant result by random variability alone. In p-value language, a “false positive” is when you say something is significant (below your p-value threshold) when it actually is not, and a “false negative” is when you say something is not significant (above the p-value threshold) when it actually is. So, in terms of multiple comparisons, if there are no adjustments made for the increased risk of false positives, this can potentially lead to inaccurate conclusions of significance.

In our example using foraging time in gray whales, we didn’t consider the context of our findings. To make this a more reliable study, we have to consider factors like the number of whales tagged (sample size!), the magnitude of noise near the tagged whales, other variables in the environment (e.g. prey availability) that could affect our results, and the ecological significance in the difference in foraging time that was found. To make robust conclusions, we need to carefully build hypotheses and study designs that will answer the questions we seek. We must then carefully choose the statistical tests that we use and explore how our data align with the assumptions that these tests make. It’s essential to contextualize our results within the bounds of our study design and broader ecological system. Finally, performing sensitivity analyses (e.g. running the same tests multiple times on slightly different datasets) ensures that our results are stable over a variety of different model parameters and assumptions. 

In the real world, there have been many studies done on the effects of noise pollution on baleen whale behavior that incorporate multiple sources of variance and bias to get robust results that show behavioral responses and physiological consequences to anthropogenic sound stressors (Melcón et al. 2012, Blair et al. 2016, Gailey et al. 2022, Lemos et al. 2022).

Moving Beyond P-values

There has been growing interest in reassessing the role of p-values in scientific inference and publishing. Scientists appreciate p-values because they provide one clear numeric threshold to determine significance of their results. However, the reality is more complicated than this binary approach. We have to explore the uncertainty around these estimates and test statistics (e.g. t-score) and what they represent ecologically. One avenue to explore might be focusing more on effect sizes and confidence intervals as more informative measures of the magnitude and precision of observed effects. There has also been a shift towards using Bayesian methods, which allow for the incorporation of prior knowledge and a more nuanced quantification of uncertainty.

Bayesian methods in particular are a leading alternative to p-values because instead of looking at how likely our observations are given a null hypothesis, we get a direct probability of the hypothesis given our data. For example, we can use Bayes factor for our noisy vs quiet gray whale behavioral t-test (Johnson et al. 2023). Bayes factor measures the likelihood of the data being observed for each hypothesis separately (instead of assuming the null hypothesis is true) so if we calculate a Bayes factor of 3 for the alternative hypothesis (HA), we could directly say that it is 3 times more likely for there to be decreased foraging time in a noisy area than for there to be no difference in the noisy vs quiet group. But that is just one example of Bayesian methods at work. The GEMM lab uses Bayesian methods in many projects from Lisa’s spatial capture-recapture models (link to blog) and Dawn’s blue whale abundance estimates (Barlow et al. 2018) to quantifying uncertainty associated with drone photogrammetry data collection methods in KC’s body size models (link to blog). 

Ultimately, the debate surrounding p-values highlights the necessity of nuanced and transparent approaches to statistical inference in scientific research. Rather than relying solely on arbitrary thresholds, researchers can consider the context, relevance, and robustness of their findings. From justifying our significance thresholds to directly describing parameters based on probability, we have increasingly powerful tools to improve the methodological rigor of our studies. 


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Baleen analyses reveals patterns in foraging ecology and stress physiology in gray whales prior to death.

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

The Eastern North Pacific (ENP) gray whale population has experienced at least two recorded Unusual Mortality Events (UMEs), from 1999–2000 and from 2019 to 2024, during which many gray whales stranded along the Pacific coast from northern Mexico to the Alaskan Arctic, USA (Martínez-Aguilar et al., 2019; Urbán, 2020). Several factors have been considered as possible causes for the high number of gray whale’s strandings, including variation in Arctic prey availability and the duration of their feeding season caused by the timing of sea ice formation and breakup (Stewart et al., 2023), starvation, anthropogenically derived toxicants, biotoxins, infectious diseases, parasites, fisheries interactions, and ship strikes (F. Gulland et al., 2005). In the most recent UME, many of the stranded whales showed signs of emaciation, indicating malnutrition as a causal factor of death (Christiansen et al., 2021; Torres et al., 2022). While the poor condition of many of the stranded whales supports the idea of starvation as a cause for these mortalities, the underlying causes of malnutrition are unknown, and it is also unclear whether the whales’ decline in body condition was rapid or gradual.

Figure 1. Gray whale with baleen exposed. Photo: GEMM Lab  NOAA/NMFS permit #16111.

Large whales face a multitude of stressors in their environment, ranging from ocean noise to contaminants, climate change, and prey shifts. Understanding how individual whales respond to these disturbances is crucial for assessing potential impacts on the population as a whole. However, monitoring the health parameters and vital rates of whales presents significant challenges due to their large size, mobility, and the vast ranges of their marine habitat. Studying stranded whales can provide valuable insights into health risks, disease susceptibility, and the impacts of pollutants and other stressors on whale populations, thus informing conservation strategies (see post). Nonetheless, the quality of information obtained from necropsies heavily relies on the timeliness of stranding reports, as decomposition begins immediately after death, limiting detailed investigations into the cause of death. Therefore, establishing a robust network capable of promptly reporting and addressing stranding events is essential (Gulland & Stockin, 2020). An effective network involves having well-trained staff, proper infrastructure, sufficient funding, and the expertise and tools necessary to gather and analyze data and samples to infer their health and causes of mortality.

During my doctoral dissertation, I worked to develop and ground truth the endocrine analyses of whale baleen as a novel sample type that can be used for retrospective assessments of the whale’s physiology (see my previous post & post). Baleen, the filter-feeding apparatus of mysticetes whales (Figure 1), consists of long fringed plates of keratinized tissue that grow continuously and slowly downward from the whale’s upper jaw. These plates are routinely collected at necropsies; and unlike other tissue types, they are durable and have minimum storage requirements; they can be preserved dry at room temperature, allowing for the analysis of both historical and current whale populations. Moreover, while most sample types used for studying whale health and physiology provide a single time-point measure of current circulating hormone levels (e.g., skin or respiratory vapor) or hold integrated information from the previous few hours or days (e.g., urine and feces), baleen tissue provides a unique opportunity for retrospective and longitudinal analyses of multiple biological parameters of the individual during the time that the tissue was grown, i.e., months to years prior to death, helping to describe the whale’s physiology, migration patterns, and exposure to pollutants (see my previous post).

In our recent study, “A longitudinal study of endocrinology and foraging ecology of subadult gray whales prior to death based on baleen analysis”, published in the journal General and Comparative Endocrinology, we examine isotope and hormone levels in the baleen of five young gray whales stranded in central Oregon during the most recent UME. Our primary objectives were to retrospectively examine the hormone and isotopic profiles of the individual whales prior to mortality, assess potential factors contributing to death, and identify the timing for the onset of chronic illness leading to mortality. Our analysis included tracing longitudinal changes in (1) stable isotope values in baleen (δ13C and δ15N), which allowed us to infer the baleen growth rate and assess the seasonal changes in diet and foraging location in large whales (Figure 2), along with the quantification of (2) two adrenal glucocorticoid steroids that are biomarkers for the whale’s stress response, (3) one thyroid hormone (triiodothyronine, T3) as an indicator of nutritional state, and (4) two sex hormones, progesterone and testosterone, to infer about reproductive status and sexual maturity. By integrating isotopic and hormonal methodologies, our study demonstrates how baleen analysis offers a comprehensive narrative of the endocrine and trophic ecology of individual whales over time.

Figure 2. Gray whales, like other large marine mammals that rely on built-up energy reserves, exhibit distinct seasonal shifts in their feeding habits. During summer, these whales feed at the ocean’s bottom, consuming organisms lower in the food chain, which is reflected in lower nitrogen values in their baleen (summer foraging). In winter, however, they must rely on their own fat reserves, causing an increase in nitrogen values (wintering). In this plot we can observe the oscillations in δ15N over time; this information allows us to estimate the baleen growth rate. Our results suggest that gray whale baleen holds a record of around 1.3 years of stable isotopes and hormone data prior to the time of death (Fernandez Ajo et al. 2024). The red cross in the X-axis, indicate the time of death. Gray whale illustration

Our endocrine assessments revealed detailed profiles of stress-related hormones (glucocorticoids, cortisol) and thyroid hormones along the lengths of the baleen. We found increased levels of cortisol in whales that died from unknown causes, starting about eight months prior to their deaths. This suggests these whales were under prolonged stress before dying. In contrast, in the case of a whale killed acutely by a killer whale, cortisol levels were low and constant prior to death, indicating this individual was likely in good health prior to the sudden attack. In terms of thyroid activity, indicated by T3 hormone levels, we found a gradual increase over several months in the whales that died of unknown causes. This pattern is not typically expected, as stress usually suppresses thyroid function. This anomaly could suggest an adaptive response to maintain body temperature and metabolism in potentially malnourished whales. Regarding the sex hormones, as expected for this age class, we found no significant fluctuations or spikes that would indicate sexual maturity in these young whales (Figure 3).

Figure 3. Longitudinal hormone profiles in an individual gray whale that died due to unknown causes (left) and one that died acutely due to orca predation (right). Note the pronounced elevations in cortisol levels (indicative of stress) and T3 prior to death in the case of unknown cause of death, while hormone levels remained low and constant prior to death in the whale acutely killed. Sex hormones do not present any clear oscillations, indicating that these whales were likely sexually immature. The red cross in the X-axis, indicate the time of death. Killer whale (Orcinus orca) illustration killer-whale

Although commercial whaling is currently banned and several whale populations show evidence of recovery, today’s whales are exposed to a variety of other human stressors that cause significant lethal and non-lethal impacts (e.g., entanglement in fishing gear, vessel strikes, shipping noise, climate change, etc.; reviewed in Thomas et al., 2016). The recovery and conservation of large whale populations is particularly important to the oceanic environment due to their key ecological role and unique biological traits (See my previous post). Our research demonstrates the strengths of using baleen as a tool for the retrospective assessments of whale endocrinology and trophic ecology. As the Eastern North Pacific gray whale population faces recurring challenges, indicated by fluctuating numbers and unusual mortality events, innovative techniques like the baleen analysis presented here, are essential to investigate the causes of mortality and inform management, helping us understand not only the immediate causes of death but also broader environmental and ecological changes affecting their survival. Broadly implementing this approach with a greater sample size of baleen collected across a larger spatial and temporal range could significantly improve our strategies for conservation and management of baleen whales.

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Christiansen, F., Rodríguez-González, F., Martínez-Aguilar, S., Urbán, J., Swartz, S., Warick, H., Vivier, F., & Bejder, L. (2021). Poor body condition associated with an unusual mortality event in gray whales. Marine Ecology Progress Series, 658, 237–252.

Gulland, F. M. D., & Stockin, K. A. (2020). Harmonizing global strandings response. European Cetacean Society Special Publication Series.

Gulland, F., Pérez-Cortés, H., Urbán, J. R., Rojas-Bracho, L., Ylitalo, G., Weir, J., Norman, S., Muto, M., Rugh, D., Kreuder, C., & Rowles, T. (2005). Eastern North Pacific gray whale (Eschrichtius robustus) unusual mortality event, 1999-2000. U.S. Department of Commerce. NOAA Technical Memorandum. NMFS-AFSC-150., March, 33 pp.

Martínez-Aguilar, S., Mariano-Meléndez, E., López-Paz, N., Castillo-Romero, F., Zaragoza-aguilar, G. A., Rivera-Rodriguez, J., Zaragoza-Aguilar, A., Swartz, S., Viloria-Gómora, L., & Urbán, J. R. (2019). Gray whale (Eschrichtius robustus) stranding records in Mexico during the winter breeding season in 2019. Report of the International Whaling Commission. Document SC/68A/CMP/14, May.

Stewart, J. D., Joyce, T. W., Durban, J. W., Calambokidis, J., Fauquier, D., Fearnbach, H., Grebmeier, J. M., Lynn, M., Manizza, M., Perryman, W. L., Tinker, M. T., & Weller, D. W. (2023). Boom-bust cycles in gray whales associated with dynamic and changing Arctic conditions. Science, 382(6667), 207–211.

Torres, L. G., Bird, C. N., 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.

Urbán, R. (2020). Gray whale stranding records in Mexico during the 2020 winter breeding season. Unpublished Paper SC/68B/CMP/13 Presented to the IWC Scientific Committee, Cambridge.

A MOSAIC of species, datasets, tools, and collaborators

By Dr. Dawn Barlow, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Imagine you are 50 nautical miles from shore, perched on the observation platform of a research vessel. The ocean is blue, calm, and seems—for all intents and purposes—empty. No birds fly overhead, nothing disturbs the rolling swells except the occasional whitecap from a light breeze. The view through your binoculars is excellent, and in the distance, you spot a disturbance at the surface of the water. As the ship gets closer, you see splashing, and a flurry of activity emerges as a large group of dolphins leap and dive, likely chasing a school of fish. They swim along with the ship, riding the bow-wave in a brief break from their activity. Birds circle in the air above them and float on the water around them. Together with your team of observers, you rush to record the species, the number of animals, their distance to the ship, and their behavior. The research vessel carries along its pre-determined trackline, and the feeding frenzy of birds and dolphins fades off behind you as quickly as it came. You return to scanning the blue water.

Craig Hayslip and Dawn Barlow scan for marine mammals from the crow’s nest (elevated observation platform) of the R/V Pacific Storm.

The marine environment is highly dynamic, and resources in the ocean are notoriously patchy. One of our main objectives in marine ecology is to understand what drives these ephemeral hotspots of species diversity and biological activity. This objective is particularly important now as the oceans warm and shift. In the context of rapid global climate change, there is a push to establish alternatives to fossil fuels that can support society’s energy needs while minimizing the carbon emissions that are a root cause of climate change. One emergent option is offshore wind, which has become a hot topic on the West Coast of the United States in recent years. The technology has the potential to supply a clean energy source, but the infrastructure could have environmental and societal impacts of its own, depending on where it is placed, how it is implemented, and when it is operational.

Northern right whale dolphins leap into the air. Photo by Craig Hayslip.

Any development in the marine environment, including alternative energy such as offshore wind, should be undertaken using the best available scientific knowledge of the ecosystem where it will be implemented. The Marine Mammal Institute’s collaborative project, Marine Offshore Species Assessments to Inform Clean energy (MOSAIC), was designed for just this reason. As the name “MOSAIC” implies, it is all about using different tools to compile different datasets to establish crucial baseline information on where marine mammals and seabirds are distributed in Oregon and Northern California, a region of interest for wind energy development.

A MOSAIC of species

The waters of Oregon and Northern California are rich with life. Numerous cetaceans are found here, from the largest species to ever live, the blue whale, to one of the smallest cetaceans, the harbor porpoise, with many species filling in the size range in between: fin whales, humpback whales, sperm whales, killer whales, Risso’s dolphins, Pacific white-sided dolphins, northern right whale dolphins, and Dall’s porpoises, to name a few. Seabirds likewise rely on these productive waters, from the large, graceful albatrosses that feature in maritime legends, to charismatic tufted puffins, to the little Leach’s storm petrels that could fit in the palm of your hand yet cover vast distances at sea. From our data collection efforts so far, we have already documented 16 cetacean species and 64 seabird species.

A Laysan albatross glides over the water’s surface. Photo by Will Kennerley.

A MOSAIC of data and tools

Schematic of the different components of the MOSAIC project. Graphic created by Solene Derville.

Through the four-year MOSAIC project, we are undertaking two years of visual surveys and passive acoustic monitoring from Cape Mendocino to the mouth of the Columbia River on the border of Oregon and Washington and seaward to the continental slope. Six comprehensive surveys for cetaceans and seabirds are being conducted aboard the R/V Pacific Storm following a carefully chosen trackline to cover a variety of habitats, including areas of interest to wind energy developers.

These dedicated surveys are complemented by additional surveys conducted aboard NOAA research vessels during collaborative expeditions in the Northern California Current, and ongoing aerial surveys in partnership with the United States Coast Guard through the GEMM Lab’s OPAL project. Three bottom-mounted hydrophones were deployed in August 2022, and are recording cetacean vocalizations and the ambient soundscape, and these recordings will be complemented by acoustic data that is being collected continuously by the Oceans Observing Initiative. In addition to these methods to collect broad-scale species distribution information, concurrent efforts are being conducted via small boats to collect individual identification photographs of baleen whales and tissue biopsy samples for genetic analysis. Building on the legacy of satellite tracking here at the Marine Mammal Institute, the MOSAIC project is breathing new life into tag data from large whales to assess movement patterns over many years and determine the amount of time spent within our study area.

A curious fin whale approaches the R/V Pacific Storm during one of the visual surveys. Photo by Craig Hayslip.
Survey tracklines extending between the Columbia River and Cape Mendocino, designed for the MOSAIC visual surveys aboard the R/V Pacific Storm.

The resulting species occurrence data from visual surveys and acoustic monitoring will be integrated to develop Species Distribution Models for the many different species in our study region. Identification photographs of individual baleen whales, DNA profiles from whale biopsy samples, and data from satellite-tagged whales will provide detailed insight into whale population structure, behavior, and site fidelity (i.e., how long they typically stay in a given area), which will add important context to the distribution data we collect through the visual surveys and acoustic monitoring. The models will be implemented to produce maps of predicted species occurrence patterns, describing when and where we expect different cetaceans and seabirds to be under different environmental conditions.

With five visual surveys down, the MOSAIC team is gearing up for one final survey this month. The hydrophones will be retrieved this summer. Then, with data in-hand, the team will dive deep into analysis.

A MOSAIC of collaborators

The MOSAIC-4 team waves from the crow’s nest (observation platform) of the R/V Pacific Storm. Photo by Craig Hayslip.

The collaborative MOSAIC team brings together a diverse set of tools. The depth of expertise here at the Marine Mammal Institute spans a broad range of disciplines, well-positioned to provide robust scientific knowledge needed to inform alternative energy development in Oregon and Northern California waters.  

I have had the pleasure of participating in three of the six surveys aboard the R/V Pacific Storm, including leading one as Chief Scientist, and have collected visual survey data aboard NOAA Ship Bell M. Shimada and from United States Coast Guard helicopters over the years that will be incorporated in the MOSAIC of datasets for the project. This ecosystem is one that I feel deeply connected to from time spent in the field. Now, I am thrilled to dive into the analysis, and will lead the modeling of the visual survey data and the integration of the different components to produce species distribution maps for cetaceans and seabirds our study region.

This project is funded by the United States Department of Energy. The Principal Investigator is the Institute’s Director Dr. Lisa Ballance, and Co-Principal Investigators include Scott Baker, Barbara Lagerquist, Rachael Orben, Daniel Palacios, Kate Stafford, and Leigh Torres of the Marine Mammal Institute; John Calambokidis of the Cascadia Research Collective; and Elizabeth Becker of ManTech International Corp. For more information, please visit the project website, and stay tuned for updates as we enter the analysis phase.

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Every breath [a whale] takes: How and why we study cetacean respiration

Clara Bird, PhD Candidate, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

We need energy to function and survive. For animals in the wild who may have limited food availability, knowing how they spend their energy is a critical question for many scientists because it fundamentally informs how we understand their decisions about where they go and what they do. The entire field of foraging theory is founded on the concept that animals optimize their ratio of energy in and energy out so that they have enough energy to survive, reproduce (pass on their genes), watch out for threats, if need be, and rest. And, if we understand an animal’s ‘typical’ energy budget, we can then try to predict how disturbance or environmental change will affect their actual energy budgets as a consequence of that change. But how do we measure energy expenditure?

The most commonly measured energy currency is oxygen. Since our cells use oxygen to produce energy (this is why we need oxygen to live), we can measure oxygen consumption as a metric of energy expenditure. The more oxygen we consume, the more energy we’re expending. In ideal lab settings, oxygen consumption can be accurately measured by placing the subject in a chamber where the oxygen flow can be controlled (Speakman, 1999). However, you can probably see how that approach is problematic for measuring oxygen consumption in most large free-living animals, especially cetaceans. It isn’t exactly feasible to put a whale in a box.

Image 1. A great tit in a metabolic chamber. Figure 1 from Broggi et al., 2009

Fortunately, a tool called a spirometer was developed to measure oxygen consumption in restrained cetaceans. A spirometer is a device that can be placed over the blowhole(s) of an individual to accurately measure the amount of air that is exhaled and inhaled (Figure 1).  For trained cetaceans in captivity (e.g., dolphins), spirometers can be used to quantify how respiration changes after the animal performs certain behaviors (Fahlman et al., 2019). The breathing patterns of diving mammals are particularly interesting because they cannot breathe during most of their exercise (energy expenditure) as they are underwater. So, their breathing patterns after a dive tell us a lot about how much energy they spent during that dive. For example, Fahlman et al. (2019) used spirometer data from dolphins in captivity to study how their breathing patterns changed while recovering from dives of different durations. Interestingly, they found that after longer dives, dolphins took larger breaths (i.e., inhaled more air) while recovering but did not change the number of breaths. This finding is particularly relevant to the work we are conducting in the GEMM lab, where we utilize breathing patterns to quantify the energy expenditure of cetaceans in the wild, where spirometers cannot be used.

Figure 1. Figure 1 from Sumich et al. (2023). Left: a spirometer being held over the blow holes of JJ, a gray whale calf at sea world in 1997; one of the rare times that a large baleen whale was in captivity and available for these measurements. Right: example of a plot created using the data from a spirometer over JJ’s blow holes. The duration of a “blow” (exhale followed by immediate inhale) is on the x-axis, the flow rate (in liters per second) is on the y-axis. The positive curve during the exhale shows that the whale strongly exhales a lot of air very quickly, then the negative curve shows the whale inhaling a lot of air very quickly.

In a previous blog, I described how inter-breath intervals (the time between consecutive blows) are useful for estimating energy expenditure in free-living cetaceans. Essentially, a shorter interval indicates that the whale was just engaged in an energetically demanding activity. When you’re recovering from a sprint, you breathe faster (i.e., with shorter inter-breath intervals), than when you’re recovering from a walk. However, a big assumption in using inter-breath intervals as a proxy for energy expenditure is that every breath is equal. But as Fahlman et al. emphasize in their 2016 paper, every blow is not equal (Fahlman et al., 2016). In addition to varying the time between breaths, an animal can vary the intensity of each breath (e.g., Fahlman et al., 2019), the duration of each breath (Sumich et al., 2023), the number of breaths, and even the expansion of their nostrils (Nazario et al., 2022; check out this blog for more).

Image 2. Gray whale blow. Source:

Altogether, this means that it’s important to measure every breath and that no one metric tells the complete story. This also means my research question focused on comparing the energetic costs of different tactics is more complicated than I originally thought. If we go back to the first blog I wrote on this topic, I was planning ons only using inter-breath intervals to estimate energy expenditure. Fast forward four years, with all my new knowledge gained on respiration variability, I’ve modified my plan and now I’m working to first understand how all these different metrics of breathing relate to each other. Then, I’ll compare how breathing varies between different foraging tactics, which is an important follow up to my questions around individual specialization of foraging tactics. If different whales are using different foraging behaviors, does that mean they’re spending different amounts of energy? If so, are certain behaviors more advantageous than others? Of course, these answers are incomplete without understanding the prey the whales are eating, but that’s something that PhD student Nat Chazal is working to understand (check out her recent blog here).  For now, I’m working on bringing integrating all the measures of breathing, then we will start putting the story together and finding some answers to our pressing questions. 

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Broggi, J., Hohtola, E., Koivula, K., Orell, M., & Nilsson, J. (2009). Long‐term repeatability of winter basal metabolic rate and mass in a wild passerine. Functional Ecology23(4), 768–773.

Fahlman, A., Brodsky, M., Miedler, S., Dennison, S., Ivančić, M., Levine, G., Rocho-Levine, J., Manley, M., Rocabert, J., & Borque-Espinosa, A. (2019). Ventilation and gas exchange before and after voluntary static surface breath-holds in clinically healthy bottlenose dolphins, Tursiops truncatus. Journal of Experimental Biology222(5), jeb192211.

Fahlman, A., van der Hoop, J., Moore, M. J., Levine, G., Rocho-Levine, J., & Brodsky, M. (2016). Estimating energetics in cetaceans from respiratory frequency: Why we need to understand physiology. Biology Open,5(4), 436–442.

Nazario, E. C., Cade, D. E., Bierlich, K. C., Czapanskiy, M. F., Goldbogen, J. A., Kahane-Rapport, S. R., Hoop, J. M. van der, Luis, M. T. S., & Friedlaender, A. S. (2022). Baleen whale inhalation variability revealed using animal-borne video tags. PeerJ10, e13724.

Speakman, J. R. (1999). The Cost of Living: Field Metabolic Rates of Small Mammals. In A. H. Fitter & D. G. Raffaelli (Eds.), Advances in Ecological Research (Vol. 30, pp. 177–297). Academic Press.

Sumich, J. L., Albertson, R., Torres, L. G., Bird, C. N., Bierlich, K. C., & Harris, C. (2023). Using audio and UAS-based video for estimating tidal lung volumes of resting and active adult gray whales (Eschrichtius robustus). Marine Mammal Science1(8).

The Dark Side of Upwelling: It’s getting harder and harder to breathe off the Oregon coast

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 depths of the productive coastal Oregon ecosystem have long held a mystery – an increasing paucity in the concentration of dissolved oxygen at depth. When dissolved oxygen concentrations dips low enough, the condition “hypoxia” can alter biogeochemical cycling in the ocean environment and threaten marine life. Essentially, organisms can’t get enough oxygen from the water, forcing them to try to escape to more favorable waters, stay and change their behavior, or suffer the consequences and potentially suffocate.

Recent work has illuminated the cause of this mysterious rise in hypoxic waters: an increase in the wind-driven oceanographic process of upwelling (Barth et al., 2024). The seasonal upwelling of cold, nutrient-rich waters underlies the incredible productivity of the Oregon coast, but its dark twin is hypoxia: when organic material in the upper layer of the water column sinks, microbial respiration processes consume dissolved oxygen in the surrounding water. In addition, the deep waters brought to the surface by upwelling are depleted in oxygen compared to the aerated surface waters. These effects combine to form an oxygen-poor water layer over the continental shelf, which typically lasts from May until October in the Northern California Current (NCC) region. The spatial extent of this layer is highly variable – hypoxic bottom waters cover 10% of the shelf in some years and up to 62% in others, presenting challenging conditions for life occupying the Oregon shelf (Peterson et al., 2013).

Figure 1. An article in The Oregonian from 2004 documents research on a hypoxia-driven “dead zone” off the Oregon coast.

While effects of hypoxia on benthic communities and some fish species are well-documented, is unclear how increasing levels of hypoxia off Oregon may impact highly mobile, migratory organisms like whales. A primary pathway is likely through their prey – particularly species that occupy hypoxic regions and depths, like the zooplankton krill. Over the continental shelf and slope, which are important krill habitat, seasonally hypoxic waters tend to extend from about 150 meters depth to the bottom. The vertical center of krill distribution in the NCC region is around 170 meters depth, suggesting that these animals encounter hypoxic conditions regularly.

Interestingly, the two main krill species off the Oregon coast, Euphausia pacifica and Thysanoessa spinifera, use different strategies to deal with hypoxic conditions. Thysanoessa spinifera krill decrease their oxygen consumption rate to better tolerate ambient hypoxia, a behavioral modification strategy called “oxyconformity”. Euphausia pacifica, on the other hand, use “oxyregulation” to maintain the same, quite high, oxygen utilization rate regardless of ambient levels – which may indicate that this species will be less able to tolerate increasingly hypoxic waters (Tremblay et al., 2020).

Figure 2. This figure from Barth et al. 2024 maps the concentration of dissolved oxygen (uM/kg; cooler colors indicate less dissolved oxygen) to show an increase in hypoxic conditions over the continental shelf and slope (green and blue colors) across seven decades in the NCC region.

Over long time scales, such environmental pressures shape species physiology, life history, and evolution. The krill species Euphausia mucronate is endemic to the Humboldt Current System off the coast of South America, which includes a region of year-round upwelling and a persistent Oxygen Minimum Zone (OMZ). Fascinatingly, Humboldt krill can live in the core of the OMZ, using metabolic adaptations that even let them survive in anoxic conditions (i.e., no oxygen in the water). Humboldt krill abundances actually increase with shallower OMZ depths and lower levels of dissolved oxygen, pointing to the huge success of this species in evolving to thrive in conditions that challenge other local krill species (Díaz-Astudillo et al., 2022).

Back home in the NCC region, will Euphausia pacifica and Thysanoessa spinifera be pressured to adapt to continually increasing levels of hypoxia? If so, will they be able to adapt? One of krill’s many superpowers is an ability to tolerate a wide range of environmental conditions, including the dramatic gradients in temperature, water density, and dissolved oxygen that they encounter during their daily vertical migrations through the water column. Both species have strategies to deal with hypoxic conditions, and this capacity has allowed them to thrive in the active upwelling region that is the NCC. Now, the question is whether increasingly hypoxic waters will eventually force a threshold that compromises the capacity of krill to adapt – and then, what will happen to these species, and the foragers dependent on them?



Barth, J. A., Pierce, S. D., Carter, B. R., Chan, F., Erofeev, A. Y., Fisher, J. L., Feely, R. A., Jacobson, K. C., Keller, A. A., Morgan, C. A., Pohl, J. E., Rasmuson, L. K., & Simon, V. (2024). Widespread and increasing near-bottom hypoxia in the coastal ocean off the United States Pacific Northwest. Scientific Reports, 14(1), 3798.

Díaz-Astudillo, M., Riquelme-Bugueño, R., Bernard, K. S., Saldías, G. S., Rivera, R., & Letelier, J. (2022). Disentangling species-specific krill responses to local oceanography and predator’s biomass: The case of the Humboldt krill and the Peruvian anchovy. Frontiers in Marine Science, 9, 979984.

Peterson, J. O., Morgan, C. A., Peterson, W. T., & Lorenzo, E. D. (2013). Seasonal and interannual variation in the extent of hypoxia in the northern California Current from 1998–2012. Limnology and Oceanography, 58(6), 2279–2292.

Tremblay, N., Hünerlage, K., & Werner, T. (2020). Hypoxia Tolerance of 10 Euphausiid Species in Relation to Vertical Temperature and Oxygen Gradients. Frontiers in Physiology, 11, 248.

How big, how blue, how beautiful! Studying the impacts of climate change on big, (and beautiful) blue whales

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

The SAPPHIRE Project is in full swing, as we spend our days aboard the R/V Star Keys searching for krill and blue whales (Figure 1) in the South Taranaki Bight (STB) region of Aotearoa New Zealand. We are investigating how changing ocean conditions impact krill availability and quality, and how this in turn impacts blue whale behavior, health, and reproduction. Understanding the link between changing environmental conditions on prey species and predators is key to understanding the larger implications of climate change on ocean food webs and each populations’ resiliency. 

Figure 1. The SAPPHIRE team searching for blue whales. Top left) KC Bierlich, top right) Dawn Barlow, bottom left) Dawn Barlow, Kim Bernard (left to right), bottom right) KC Bierlich, Dawn Barlow, Leigh Torres, Mike Ogle (left to right).  

One of the many components of the SAPPHIRE Project is to understand how foraging success of blue whales is influenced by environmental variation (see this recent blog written by Dr. Dawn Barlow introducing each component of the project). When you cannot go to a grocery store or restaurant any time you are hungry, you must rely on stored energy from previous feeds to fuel energy needs. Body condition reflects an individual’s stored energy in the body as a result of feeding and thus represents the foraging success of an individual, which can then affect its potential for reproductive output and the individual’s overall health (see this previous blog). As discussed in a previous blog, drones serve as a valuable tool for obtaining morphological measurements of whales to estimate their body condition. We are using drones to collect aerial imagery of pygmy blue whales to obtain body condition measurements late in the foraging season between years 2024 and 2026 of the SAPPHIRE Project (Figure 2). We are quantifying body condition as Body Area Index (BAI), which is a relative measure standardized by the total length of the whale and well suited for comparing individuals and populations (Figure 3). 

The GEMM Lab recently published an article led by Dr. Dawn Barlow where we investigated the differences in BAI between three blue whale populations: Eastern North Pacific blue whales feeding in Monterey Bay, California; Chilean blue whales feeding in the Corcovado Gulf; and New Zealand Pygmy blue whales feeding in the STB (Barlow et al., 2023). These three populations are interesting to compare since blue whales that feed in Monterey Bay and Corcovado Gulf migrate to and from these seasonally productive feeding grounds, while the Pygmy blue whales stay in Aotearoa New Zealand year-round. Interestingly, the Pygmy blue whales had higher BAI (were fatter) compared to the other two regions despite relatively lower productivity in their foraging grounds. This difference in body condition may be due to different life history strategies where the non-migratory Pygmy blue whales may be able to feed as opportunities arrive, while the migratory strategies of the Eastern North Pacific and Chilean blue whales require good timing to access high abundant prey. Another interesting and unexpected result from our blue whale comparison was that Pygmy blue whales are not so “pygmy”; they are actually the same size as Eastern North Pacific and Chilean blue whales, with an average size around 22 m. Our findings from this blue whale comparison leads us to more questions about how environmental conditions that vary from year to year influence body condition and reproduction of these “not so pygmy” blue whales. 

Figure 2. An aerial image of a Pygmy blue whale in the South Taranaki Bight region of Aotearoa New Zealand collected during the SAPPHIRE 2024 field season using a DJI Inspire 2 drone. 
Figure 3. A drone image of a Pygmy blue whale and the length and body width measurements used to estimate Body Area Index (BAI), represented by the shaded blue region. Width measurements will also be used to help identify pregnant individuals.

The GEMM Lab has been studying this population of Pygmy blue whales in the STB since 2013 and found that years designated as a marine heatwave resulted with a reduction in blue whale feeding activity. Interestingly, breeding activity is also reduced during marine heatwaves in the following season when compared to the breeding season following a more productive, typical foraging season. These findings indicate that fluctuations in the environment, such as marine heatwaves, may affect not only foraging success, but also reproduction in Pygmy blue whales. 

To help us better understand reproductive patterns across years, we will use body width measurements from drone images paired with hormone concentrations collected from fecal and biopsy samples to identify pregnant individuals. Progesterone is a hormone secreted in the ovaries of mammals during the estrous cycle and gestation, making it the predominant hormone responsible for sustaining pregnancy. Recently, the GEMM Lab’s Dr. Alejandro Fernandez-Ajo wrote a blog discussing his publication identifying pregnant individual gray whales using drone-based body width measurements and progesterone concentrations from fecal samples (Fernandez et al., 2023). While individuals that were pregnant had higher levels of progesterone compared to when they were not pregnant, the body width at 50% of the body length served as a more reliable method for detecting pregnancy in gray whales. We will use similar methods to help identify pregnancy in Pygmy blue whales for the SAPPHIRE Project where will we examine body width measurement paired with progesterone concentrations collected from fecal and biopsy samples to identify pregnant individuals. We hope our work will help to better understand how climate change will influence Pygmy blue whale body condition and reproduction, and thus the overall health and resiliency of the population. Stay tuned! 


Barlow, D. R., Bierlich, K. C., Oestreich, W. K., Chiang, G., Durban, J. W., Goldbogen, J. A., Johnston, D. W., Leslie, M. S., Moore, M. J., Ryan, J. P., & Torres, L. G. (2023). Shaped by Their Environment: Variation in Blue Whale Morphology across Three Productive Coastal Ecosystems. Integrative Organismal Biology, 5(1).

Fernandez Ajó, A., Pirotta, E., Bierlich, K. C., Hildebrand, L., Bird, C. N., Hunt, K. E., Buck, C. L., New, L., Dillon, D., & Torres, L. G. (2023). Assessment of a non-invasive approach to pregnancy diagnosis in gray whales through drone-based photogrammetry and faecal hormone analysis. Royal Society Open Science10(7), 230452.