Who, where, when: Estimating individual space use patterns of PCFG gray whales

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

Understanding how baleen whales are affected by human activity is a central goal for many research projects in the GEMM Lab. The overarching goal of the GRANITE (Gray whale Response to Ambient Noise Informed by Technology and Ecology) project is to quantify baleen whale physiological response to different stressors (e.g., boat presence and noise) and model the subsequent impacts of these stressors on the population. We will achieve this goal by implementing our long-term, replicate dataset of Pacific Coast Feeding Group (PCFG) gray whales into a framework called population consequences of disturbance (PCoD). I will not go into the details of PCoD in this blog (but I wrote a post a few years ago that you can revisit). Instead, I will explain the approach I am taking to assess where and when individual whales spend time in our study area, which will form an essential component of PCoD and be one of the chapters of my PhD dissertation.

Individuals in a population are unlikely to be exposed to a stressor in a uniform way because they make decisions differently based on intrinsic (e.g., sex, age, reproductive status) and extrinsic (e.g., environment, prey, predators) factors (Erlinge & Sandell 1986). For example, a foraging female gray whale who is still nursing a calf will need to consider factors that are different to ones that an adult single male might need to consider when choosing a location to feed. These differences in decision-making exist across the whole population, which makes it important to understand where individuals are spending time and how they overlap with stressors in space and time before trying to quantify the impacts of stressors on the population as a whole (Pirotta et al. 2018). I am currently working on an analysis that will determine an individual’s exposure to a number of stressors based on their space use patterns. 

We can monitor space use patterns of individuals in a population through time using spatial capture-recapture techniques. As the name implies, a spatial capture-recapture technique involves capturing an individual in a marked location during a sampling period, releasing it back into the population, and then (hopefully) re-capturing it during another sampling period in the future, at either the same or a different location. With enough repeat sampling events, the method should build spatial capture histories of individuals through time to better understand an individual’s space use patterns (Borchers & Efford 2008). While the use of the word capture implies that the animal is being physically caught, this is not necessarily the case. Individuals can be “captured” in a number of non-invasive ways, including by being photographed, which is how we “capture” individual PCFG gray whales. These capture-recapture methods were first pioneered in terrestrial systems, where camera traps (i.e., cameras that take photos or videos when a motion sensor is triggered) are set up in a systematic grid across a study area (Figure 1; Royle et al. 2009, Gray 2018). Placing the cameras in a grid system ensures that there is an equal distribution of cameras throughout the study area, which means that an animal theoretically has a uniform chance of being captured. However, because we know that individuals within a population make space use decisions differently, we assume that individuals will distribute themselves differently across a landscape, which will manifest as individuals having different centers of their spatial activity. The probability of capturing an individual is highest when a camera trap is at that individual’s activity center, and the cameras furthest away from the individual’s activity center will have the lowest probability of capturing that individual (Efford 2004). By using this principle of probability, the data generated from spatial capture-recapture field methods can be modelled to estimate the activity centers and ranges for all individuals in a population. The overlap of an individual’s activity center and range can then be compared to the spatiotemporal distribution of stressors that an individual may be exposed to, allowing us to determine whether and how an individual has been exposed to each stressor. 

Figure 1. Example of camera trap grid in a study area. Figure taken from Gray (2018).

While capture-recapture methods were first developed in terrestrial systems, they have been adapted for application to marine populations, which is what I am doing for our GRANITE dataset of PCFG gray whales. Together with a team of committee members and GRANITE collaborators, I am developing a Bayesian spatial capture-recapture model to estimate individual space use patterns. In order to mimic the camera trap grid system, we have divided our central Oregon coast study area into latitudinal bins that are approximately 1 km long. Unfortunately, we do not have motion sensor activated cameras that automatically take photographs of gray whales in each of these latitudinal bins. Instead, we have eight years of boat-based survey effort with whale encounters where we collect photographs of many individual whales. However, as you now know, being able to calculate the probability of detection is important for estimating an individual’s activity center and range. Therefore, we calculated our spatial survey effort per latitudinal bin in each study year to account for our probability of detecting whales (i.e., the area of ocean in km2 that we surveyed). Next, we tallied up the number of times we observed every individual PCFG whale in each of those latitudinal bins per year, thus creating individual spatial capture histories for the population. Finally, using just those two data sets (the individual whale capture histories and our survey effort), we can build models to test a number of different hypotheses about individual gray whale space use patterns. There are many hypotheses that I want to test (and therefore many models that I need to run), with increasing complexity, but I will explain one here.

Over eight years of field work for the GRANITE project, consisting of over 40,000 km2 of ocean surveyed with 2,169 sightings of gray whales, our observations lead us to hypothesize that there are two broad space use strategies that whales use to optimize how they find enough prey to meet their energetic needs. For the moment, we are calling these strategies ‘home-body’ and ‘roamer’. As the name implies, a home-body is an individual that stays in a relatively small area and searches for food in this area consistently through time. A roamer, on the other hand, is an individual that travels and searches over a greater spatial area to find good pockets of food and does not generally tend to stay in just one place. In other words, we except a home-body to have a consistent activity center through time and a small activity range, while a roamer will have a much larger activity range and its activity center may vary more throughout the years (Figure 2). 

Figure 2. Schematic representing one of the hypotheses we will be testing with our Bayesian spatial capture-recapture models. The schematic shows the activity centers (the circles) and activity ranges (vertical lines attached to the circles) of two individuals (green and orange) across three years in our central Oregon study area. The green individual represents our hypothesized idea of a home-body, whereas the orange individuals represents our idea of a roamer.

While this hypothesis sounds straightforward, there are a lot of decisions that I need to make in the Bayesian modeling process that can ultimately impact the results. For example, do all home-bodies in a population have the same size activity range or can the size vary between different home-bodies? If it can vary, by how much can it vary? These same questions apply for the roamers too. I have a long list of questions just like these, which means a lot of decision-making on my part, and that long list of hypotheses I previously mentioned. Luckily, I have a fantastic team made up of Leigh, committee members, and GRANITE collaborators that are guiding me through this process. In just a few more months, I hope to reveal how PCFG individuals distribute themselves in space and time throughout our central Oregon study area, and hence describe their exposure to different stressors. Stay tuned! 

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Borchers DL, Efford MG (2008) Spatially explicit maximum likelihood methods for capture-recapture studies. Biometrics 64:377-385.

Efford M (2004) Density estimation in live-trapping studies. Oikos 106:598-610.

Erlinge S, Sandell M (1986) Seasonal changes in the social organization of male stoats, Mustela erminea: An effect of shifts between two decisive resources. Oikos 47:57-62.

Gray TNE (2018) Monitoring tropical forest ungulates using camera-trap data. Journal of Zoology 305:173-179.

Pirotta E, Booth CG, Costa DP, Fleishman E, Kraus SD, and others (2018) Understanding the population consequences of disturbance. Ecology and Evolution 8(19):9934–9946.Royle J, Nichols J, Karanth KU, Gopalaswamy AM (2009) A hierarchical model for estimating density in camera-trap studies. Journal of Applied Ecology 46:118-127.

Learning from the unexpected: the first field season of the SAPPHIRE project

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

The SAPPHIRE project’s inaugural 2024 field season has officially wrapped up, and the team is back on shore after an unexpected but ultimately fruitful research cruise. The project aims to understand the impacts of climate change on blue whales and krill, by investigating their health under variable environmental conditions. In order to assess their health, however, a crucial first step is required: finding krill, and finding whales. The South Taranaki Bight (STB) is a known foraging ground where blue whales typically feed on krill found in the cool and productive upwelled waters. This year, however, both krill and blue whales were notoriously absent from the STB, leaving us puzzled as we compulsively searched the region in between periods of unworkable weather (including an aerial survey one afternoon).

A map of our survey effort during the 2024 field season. Gray lines represent our visual survey tracklines, with the aerial survey shown in the dashed line. Red points show blue whale sighting locations. Purple stars are the deployment locations of two hydrophones, which will record over the next year.

The tables felt like they were turning when we finally found a blue whale off the west coast of the South Island, and were able to successfully fly the drone to collect body condition information, and collect a fecal sample for genetic and hormone analysis. Then, we returned to the same pattern. Days of waiting for a weather window in between fierce winds, alternating with days of searching and searching, with no blue whales or krill to be found. Photogrammetry measurements of our drone data over the one blue whale we found determined it to be quite small (only ~17 m) and in poor body condition. The only krill we were able to find and collect were small and sparsely mixed in to a massive gelatinous swarm of salps. Where were the whales? Where was their prey?

Above: KC Bierlich and Dawn Barlow search for blue whales. Below: salps swarm beneath the surface.

Then, a turn of events. A news story with the headline “Acres of krill washing up on the coastline” made its way to our inboxes and news feeds. The location? Kaikoura. On the other side of the Cook Strait, along the east coast of the South Island. With good survey coverage in the STB resulting in essentially no appearances of our study species, this report of krill presence along with a workable weather forecast in the Kaikoura area had our attention. In a flurry of quick decision-making (Leigh to Captain: “Can we physically get there?” Captain to Leigh: “Yes, we can.” Leigh to Captain: “Let’s go.”), we turned the vessel around and surfed the swells to the southeast at high speed.

The team in action aboard the R/V Star Keys, our home for the duration of the three-week survey.

Twelve hours later we arrived at dusk and anchored off the small town of Kaikoura, with plans to conduct a net tow for krill before dawn the next morning. But the krill came to us! In the wee hours of the morning, the research vessel was surrounded by swarming krill. The dense aggregation made the water appear soup-like, and attracted a school of hungry barracuda. These abundant krill were just what was needed to run respiration experiments on the deck, and to collect samples to analyze their calories, proteins, and lipids back in the lab.

Left: An illuminated swarm of krill just below the surface. Right: A blue whale comes up for air with an extended buccal pouch, indicating a recent mouthful of krill. Drone piloted by KC Bierlich.

With krill in the area, we were anxious to find their blue whale predators, too. Once we began our visual survey effort, we were alerted by local whale watchers of a blue whale sighting. We headed straight to this location and got to work. The day that followed featured another round of krill experiments, and a few more blue whale sightings. Predator and prey were both present, a stark contrast to our experience in the previous weeks within the STB and along the west coast of the South Island. The science team and crew of the R/V Star Keys fell right into gear, carefully maneuvering around these ocean giants to collect identification photos, drone flights, and fecal samples, finding our rhythm in what we came here to do. We are deeply grateful to the regional managers, local Iwi representatives, researchers, and tourism operators that supported making our time in Kaikoura so fruitful, on just a moment’s notice.

The SAPPHIRE 2024 field team on a day of successful blue whale sightings. Clockwise, starting top left: Dawn Barlow and Leigh Torres following a sunset blue whale sighting, Mike Ogle in position for biopsy sample collection, Kim Bernard collecting blue whale dive times, KC Bierlich collecting identification photos.

What does it all mean? It’s hard to say right now, but time and data analysis will hopefully tell. While this field season was certainly unexpected, it was valuable in many ways. Our experiences this year emphasize the pay-off of being adaptable in the field to maximize time, money, and data collection efforts (during our three-week cruise we slept in 10 different ports or anchorages, did an aerial survey, and rapidly changed our planned study area). Oftentimes, the cases that initially “don’t make sense” are the ones that end up providing key insights into larger patterns. No doubt this was a challenging and at times frustrating field season, but it could also be the year that provides the greatest insights. After two more years of data collection, it will be fascinating to compare this year’s blue whale and krill data in the greater context of environmental variability.

A blue whale comes up for air. Photo by Dawn Barlow.

One thing is clear, the oceans are without question already experiencing the impacts of global climate change. This year solidified the importance of our research, emphasizing the need to understand how krill—a crucial marine prey item—and their predators are being affected by warming and shifting oceans.  

A blue whale at sunset, off Kaikoura. Photo by Leigh Torres.

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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). https://doi.org/10.1093/iob/obad039

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. https://doi.org/10.1098/rsos.230452

Phases and Feelings of the Scientific Journey

Leigh Torres, Associate Professor, PI of the GEMM Lab

There are many phases of a scientific journey, which generally follows a linear path (although I recognize that the process is certainly iterative at times to improve and refine). The scientific journey typically starts with an idea or question, bred from curiosity and passion. The journey hopefully ends with new knowledge, a useful application (e.g., tool or management outcome), and more questions in need of answers, providing a sense of success and pride. But along this path, there are many more phases, with many more emotions. As we begin the four-year SAPPHIRE project, I have already experienced a range of emotions, and I am certain more will come my way as I again wander through the many phases and feeling of science:

Generation of idea or questionCuriosity, passion, wonder
Build the team and develop the funding proposalDrive, dreaming big, team management, belief in the importance of your proposed work
Notice of funding proposal successDisbelief, excitement, and pride, followed quickly by feeling daunted, and self-doubt about the ability to pull off what you said you would do.
*Prep for fieldwork/experiment/data collectionFrantic and overwhelmed by the need to remember all the details that make or break the research; lists, lists, lists; pressure to get organized and stay within your budget. Anticipation, exhaustion.
*Outreach/Engagement/CommunicationEagerness to share and connect; Pressure to build relationships and trust; make sure the research is meaningful and accessible to local communities
*Fieldwork/experiment/data collection/data analysisSigh of relief to be underway, accompanied by big pressure to achieve: gotta do what you said you would do.
Preparation of scientific publications and reportsExcitement for data synthesis: What will the results say? What are the answers to your burning questions? Were your hypotheses correct? With a good dose of apprehension of peer feedback and critical reviews.
Publications and reportsSatisfaction to see outputs and results from hard work being broadly disseminated.
Project end with final reportFeeling of great accomplishment, but now need to develop the next project and get the funding… the cycle continues.

*After months of intense preparation for our field research component of the SAPPHIRE project in Aotearoa New Zealand (permits, equipment purchasing, community engagement, gathering supplies, learning how to use new equipment, vessel contracting, overseas shipping, travel arrangements, vessel mobilization, oh the list goes on!), we have just stepped off the vessel after 3 full days collecting data. I have cycled through all these emotions many times, and now I feel both exhausted and elated. We are implementing our plan, and we now have data in-hand. Worry creeps in all the time: we need to do more, do better. But I also know that our team is excellent and with patience, blessings from the weather gods, and our continued hard work, we will succeed, learn, and share. As SAPPHIRE chargers ahead to understand the impacts of climate change on marine prey (krill) and predators (blue whales), I am ready for the continued mix of emotions that comes with science.

Photo montage of our awesome SAPPHIRE team in prep mode and during data collection in the South Taranaki Bight within Aotearoa New Zealand.

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Blue whales, krill, and climate change: introducing the SAPPHIRE project

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

The world is warming. Ocean ecosystems are experiencing significant and rapid impacts of climate change. However, the cascading effects on marine life are largely unknown. Thus, it is critical to understand how – not just if – environmental change impacts the availability and quality of key prey species in ocean food webs, and how these changes will impact marine predator health and population resilience. With these pressing knowledge gaps in mind, we are thrilled to launch a new project “Marine predator and prey response to climate change: Synthesis of Acoustics, Physiology, Prey, and Habitat in a Rapidly changing Environment (SAPPHIRE).”  We will examine how changing ocean conditions affect the availability and quality of krill, and thus impact blue whale behavior, health, and reproduction. This large-scale research effort is made possible with funding from the National Science Foundation.

The SAPPHIRE project takes place in the South Taranaki Bight (STB) region of Aotearoa New Zealand, and before diving into our new research plans, let’s reflect briefly on what we know so far about this study system based on our previous research. Our collaborative research team has studied blue whales in the STB since 2013 to document the population, understand their ecology and habitat use, and inform conservation management. We conducted boat-based surveys and used hydrophones to record the underwater soundscape, and found the following:

  • Blue whales in Aotearoa New Zealand are a unique population, genetically distinct from all other known populations in the Southern Hemisphere, with an estimated population size of 718 (95% CI = 279 – 1926).1
  • Blue whales reside in the STB region year-round, with feeding and breeding vocalizations detected nearly every day of the year.2,3
  • Wind-driven upwelling over Kahurangi shoals moves a plume of cold, nutrient-rich waters into the STB, supporting aggregations of krill, and thereby critical feeding opportunities for blue whales in spring and summer.4–6
  • We developed predictive models to forecast blue whale distribution up to three weeks in advance, providing managers with a real-time tool in the form of a desktop application to produce daily forecast maps for dynamic management.7
  • During marine heatwaves, blue whale feeding activity was substantially reduced in the STB. Interestingly, their breeding activity was also reduced in the following season when compared to the breeding season following a more productive, typical foraging season. This finding indicates that shifting environmental conditions, such as marine heatwaves and climate change, may have consequences to not just foraging success, but the population’s reproductive patterns.3
A blue whale comes up for air in the South Taranaki Bight. Photo by Leigh Torres.

Project goals

Building on this existing knowledge, we aim to gain understanding of the health impacts of environmental change on krill and blue whales, which can in turn inform management decisions. Over the next three years (2024-2026) we will use multidisciplinary methods to collect data in the field that will enable us to tackle these important but challenging goals. Our broad objectives are to:

  1. Assess variation in krill quality and availability relative to rising temperatures and different ocean conditions,
  2. Document how blue whale body condition and hormone profiles change relative to variable environmental and prey conditions,
  3. Understand how environmental conditions impact blue whale foraging and reproductive behavior, and
  4. Integrate these components to develop novel Species Health Models to predict predator and prey whale population response to rapid environmental change.

Kicking off fieldwork

This coming January, we will set sail aboard the R/V Star Keys and head out in search of blue whales and krill in the STB! Five of our team members will spend three weeks at sea, during which time we will conduct surveys for blue whale occurrence paired with active acoustic assessment of krill availability, fly Unoccupied Aircraft Systems (UAS; “drones”) over whales to determine body condition and potential pregnancy, collect tissue biopsy samples to quantify stress and reproductive hormone levels, deploy hydrophones to record rates of foraging and reproductive calls by blue whales, and conduct on-board controlled experiments on krill to assess their response to elevated temperature.

The team in action aboard the R/V Star Keys in February 2017. Photo by L. Torres.

The moving pieces are many as we work to obtain research permits, engage in important consultation with iwi (indigenous Māori groups), procure specialized scientific equipment, and make travel and shipping arrangements. The to-do lists seem to grow just as fast as we can check items off; such is the nature of coordinating an international, multidisciplinary field effort. But it will pay off when we are underway, and I can barely contain my excitement to back on the water with this research team.

Our team has not collected data in the STB since 2017. We know so much more now than we did when studies of this blue whale population were just beginning. For example, we are eager to put our blue whale forecast tool to use, which will hopefully enable us to direct survey effort toward areas of higher blue whale density to maximize data collection. We are keen to see what new insights we gain, and what new questions and challenges arise.

Research team

The SAPPHIRE project will only be possible with the expertise and coordination of the many members of our collaborative group. We are all thrilled to begin this research journey together, and eager to share what we learn.

Principal Investigators:

Research partners and key collaborators:

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1.          Barlow DR, Torres LG, Hodge KB, Steel D, Baker CS, Chandler TE, Bott N, Constantine R, Double MC, Gill P, Glasgow D, Hamner RM, Lilley C, Ogle M, Olson PA, Peters C, Stockin KA, Tessaglia-Hymes CT, Klinck H. Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger Species Res. 2018;36:27–40.

2.          Barlow DR, Klinck H, Ponirakis D, Holt Colberg M, Torres LG. Temporal occurrence of three blue whale populations in New Zealand waters from passive acoustic monitoring. J Mammal. 2022;

3.          Barlow DR, Klinck H, Ponirakis D, Branch TA, Torres LG. Environmental conditions and marine heatwaves influence blue whale foraging and reproductive effort. Ecol Evol. 2023;13:e9770.

4.          Barlow DR, Klinck H, Ponirakis D, Garvey C, Torres LG. Temporal and spatial lags between wind, coastal upwelling, and blue whale occurrence. Sci Rep. 2021;11(6915):1–10.

5.          Barlow DR, Bernard KS, Escobar-Flores P, Palacios DM, Torres LG. Links in the trophic chain: Modeling functional relationships between in situ oceanography, krill, and blue whale distribution under different oceanographic regimes. Mar Ecol Prog Ser. 2020;642:207–25.

6.          Torres LG, Barlow DR, Chandler TE, Burnett JD. Insight into the kinematics of blue whale surface foraging through drone observations and prey data. PeerJ. 2020;8:e8906.

7.          Barlow DR, Torres LG. Planning ahead: Dynamic models forecast blue whale distribution with applications for spatial management. J Appl Ecol. 2021;58(11):2493–504.

Familiar flukes and flanks: The 9th GRANITE field season is underway

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

The winds are consistently (and sometimes aggressively) blowing from the north here on the Oregon coast, which can only mean one thing – summer has arrived! Since mid-May, the GRANITE (Gray whale Response to Ambient Noise Informed by Technology and Ecology) team has been looking for good weather windows to survey for gray whales and we have managed to get five great field work days already. In today’s blog post, I am going to share what (and who) we have seen so far.

On our first day of the field season, PI Leigh Torres, postdoc KC Bierlich and myself, were joined by a special guest: Dr. Andy Read. Andy is the director of the Duke University Marine Lab, where he also runs his own lab, which focuses on conservation biology and ecology of marine vertebrates. Andy was visiting the Hatfield Marine Science Center as part of the Lavern Weber Visiting Scientist program and was hosted here by Leigh. For those of you that do not know, Andy was Leigh’s graduate school advisor at Duke where she completed her Master’s and doctoral degrees. It felt very special to have Andy on board our RHIB Ruby for the day and to introduce him to some friends of ours. The first whale we encountered that day was “Pacman”. While we are always excited to re-sight an individual that we know, this sighting was especially mind-blowing given the fact that Leigh had “just” seen Pacman approximately two months earlier in Guerrero Negro, one of the gray whale breeding lagoons in Mexico (read this blog about Leigh and Clara’s pilot project there). Aside from Pacman, we saw five other individuals, all of which we had seen during last year’s field season. 

The first day of field work for the 2023 GRANITE field season! From left to right: Leigh Torres, Lisa Hildebrand, Andy Read, and KC Bierlich. Source: L. Torres.

Since that first day on the water, we have conducted field work on four additional days and so far, we have only encountered known individuals in our catalog. This fact is exciting because it highlights the strong site fidelity that Pacific Coast Feeding Group (PCFG) gray whales have to areas within their feeding range. In fact, I am examining the residency and space use of each individual whale we have observed in our GRANITE study for one of my PhD chapters to better understand the level of fidelity individuals have to the central Oregon coast. Furthermore, this site fidelity underpins the unique, replicate data set on individual gray whale health and ecology that the GRANITE project has been able to progressively build over the years. So far during this field season in 2023, we have seen 13 unique individuals, flown the drone over 10 of them and collected four fecal samples from two, which represent critical data points from early on in the feeding season.

Our sightings this year have not only highlighted the high site fidelity of whales to our study area but have also demonstrated the potential for internal recruitment of calves born to “PCFG mothers” into the PCFG. Recruitment to a population can occur in two ways: externally (individuals immigrate into a population from another population) or internally (calves born to females that are part of the population return to, or stay, within their mothers’ population). Three of the whales we have seen so far this year are documented calves from females that are known to consistently use the PCFG range, including our central Oregon coast study area. In fact, we documented one of these calves, “Lunita”, just last year with her mother (see Clara’s recap of the 2022 field season blog for more about Lunita). The average calf survival estimate between 1997-2017 for the PCFG was 0.55 (Calambokidis et al. 2019), though it varied annually and widely (range: 0.34-0.94). Considering that there have been years with calf survival estimates as low as ~30%, it is therefore all the more exciting when we re-sight a documented calf, alive and well!

“Lunita”, an example of successful internal recruitment

We have also been collecting data on the habitat and prey in our study system by deploying our paired GoPro/RBR sensor system. We use the GoPro to monitor the benthic substrate type and relative prey densities in areas where whales are feeding. The RBR sensor collects high-frequency, in-situ dissolved oxygen and temperature data, enabling us to relate environmental metrics to relative prey measurements. Furthermore, we also collect zooplankton samples with a net to assess prey community and quality. On our five field work days this year, we have predominantly collected mysid shrimp, including gravid (a.k.a. pregnant) individuals, however we have also caught some Dungeness and porcelain crab larvae. The GEMM Lab is also continuing our collaboration with Dr. Susanne Brander’s lab at OSU and her PhD student Lauren Kashiwabara, who plan on conducting microplastic lab experiments on wild-caught mysid shrimp. Their plan is to investigate the growth rates of mysid shrimp under different temperature, dissolved oxygen, and microplastic load conditions. However, before they can begin their experiments, they need to successfully culture the mysids in the lab, which is why we collect samples for them to use as their ‘starter culture’. Stay tuned to hear more about this project as it develops!

So, all in all, it has been an incredibly successful start to our field season, marked by the return of many familiar flukes and flanks! We are excited to continue collecting rock solid GRANITE data this summer to increase our efforts to understand gray whale ecology and physiology. 



Calambokidis, J., Laake, J., and Perez, A. (2019). Updated analyses of abundance and population structure of seasonal gray whales in the Pacific Northwest, 1996-2017. IWC, SC/A17/GW/05 for the Workshop on the Status of North Pacific Gray Whales. La Jolla: IWC.

New GEMM Lab publication reveals how blue whale feeding and reproductive effort are related to environmental conditions

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

Learning by listening

Studying mobile marine animals that are only fleetingly visible from the water’s surface is challenging. However, many species including baleen whales rely on sound as a primary form of communication, producing different vocalizations related to their fundamental needs to feed and reproduce. Therefore, we can learn a lot about these elusive animals by monitoring the patterns of their calls. In the final chapter of my PhD, we set out to study blue whale ecology and life history by listening. I am excited to share our findings, recently published in Ecology and Evolution.

Blue whales produce two distinct types of vocalizations: song is produced by males and is hypothesized to play a role in breeding behavior, and D calls are a hypothesized social call produced by both sexes in association with feeding behavior. We analyzed how these different calls varied seasonally, and how they related to environmental conditions.

This paper is a collaborative study co-authored by Dr. Holger Klinck and Dimitri Ponirakis of the K. Lisa Yang Center for Conservation Bioacoustics, Dr. Trevor Branch of the University of Washington, and GEMM Lab PI Dr. Leigh Torres, and brings together multiple methods and data sources. Our findings shed light on blue whale habitat use patterns, and how climate change may impact both feeding and reproduction for this species of conservation concern.

The South Taranaki Bight: an ideal study system

Baleen whales typically migrate between high-latitude, productive feeding grounds and low-latitude breeding grounds. However, the New Zealand blue whale population is present in the South Taranaki Bight (STB) region year-round, which uniquely enabled us to monitor their behavior, ecology, and life history across seasons and years from a single location. We recorded blue whale vocalizations from Marine Autonomous Recording Units (MARUs) deployed at five locations in the STB for two full years (Fig. 1).

Figure 1. Study area map and blue whale call spectrograms. Left panel: map of the study area in the South Taranaki Bight region, with hydrophone (marine autonomous recording unit; MARU) locations denoted by the stars. Gray lines show bathymetry contours at 50 m depth increments, from 0 to 500 m. Location of the study area within New Zealand is indicated by the inset map. Right panels: example spectrograms of the two blue whale call types examined: the New Zealand song recorded on 31 May 2016 (top) and D calls recorded 20 September 2016 (bottom). Figure reproduced from Barlow et al. (2023).

We found that the two vocalization types had different seasonal occurrence patterns (Fig. 2). D calls were associated with upwelling conditions that indicate feeding opportunities, lending evidence for their function as a foraging-related call.

Figure 2. Average annual cycle in the song intensity index (dark blue) and D calls (green) per day of the year, computed across all hydrophone locations and the entire two-year recording period. Figure reproduced from Barlow et al. (2023).

In contrast, blue whale song showed a very clear seasonal peak in the fall and was less obviously correlated with environmental conditions. To investigate the hypothesized function of song as a breeding call, we turned to a perhaps unintuitive source of information: historical whaling records. Whenever a pregnant whale was killed during commercial whaling operations, the length of the fetus was measured. By looking at the seasonal pattern in these fetal lengths, we can presume that births occur around the time of year when fetal lengths are at their longest. The records indicated April-May. By back-calculating the 11-month gestation time for a blue whale, we can presume that mating occurs generally in May-June, which is the exact time of the peak in song intensity from our recordings (Fig. 3).

Figure 3. Annual song intensity and the breeding cycle. Top panel: average yearly cycle in song intensity index, computed across the five hydrophone locations and the entire recording period; dark blue line represents a loess smoothed fit. Bottom panel: fetal length measurements from whaling catch records for Antarctic blue whales (gray, measurements rounded to the nearest foot), pygmy blue whales in the southern hemisphere (blue, measurements rounded to the nearest centimeter). Measurements from blue whales caught within the established range of the New Zealand population are denoted by the dark red triangles. Calving presumably takes place around or shortly after fetal lengths are at their maximum (April–May), which implies that mating likely occurs around May–June, coincident with the peak song intensity. Figure reproduced from Barlow et al. (2023).

With this evidence for D calls as feeding-related calls and song as breeding-related calls, we had a host of new questions, we used this gained knowledge to explore how changing environmental conditions might impact multiple life history processes for New Zealand blue whales

Marine heatwaves impact multiple life history processes

Our study period between January 2016 and February 2018 spanned both typical upwelling conditions and dramatic marine heatwaves in the STB region. While we previously documented that the marine heatwave of 2016 affected blue whale distribution, the population-level impacts on feeding and reproductive effort remained unknown. In our recent study, we found that during marine heatwaves, D calls were dramatically reduced compared to during productive upwelling conditions. During the fall breeding peak, song intensity was likewise dramatically reduced following the marine heatwave. This relationship indicates that following poor feeding conditions, blue whales may invest less effort in reproduction. As marine heatwaves are projected to become more frequent and more intense under global climate change, our findings are perhaps a warning for what is to come as animal populations must contend with changing ocean conditions.

More than a decade of research on New Zealand blue whales

Ten years ago, Leigh first put forward a hypothesis that the STB region was an undocumented blue whale foraging ground based on multiple lines of evidence (Torres 2013). Despite pushback and numerous challenges, Leigh set out to prove her hypothesis through a comprehensive, multi-year data collection effort. I was lucky enough to join the team in 2016, first as a Masters’ student, and then as a PhD student. In the time since Leigh’s hypothesis, we not only documented the New Zealand blue whale population (Barlow et al. 2018), we learned a great deal about what drives blue whale feeding behavior (Torres et al. 2020) and habitat use patterns (Barlow et al. 2020, 2021), and developed forecast models to predict blue whale distribution for dynamic management of the STB (Barlow & Torres 2021). We also documented their unique, year-round presence in the STB, distinct from the migratory or vagrant presence of other blue whale populations (Barlow et al. 2022b). We now understand how marine heatwaves impact both feeding opportunities and reproductive effort (Barlow et al. 2023). We even analyzed blue whale skin condition (Barlow et al. 2019) and acoustic response to earthquakes (Barlow et al. 2022a) along the way. A decade later, it is humbling to reflect on how much we have learned about these whales. This paper is also the final chapter of my PhD, and as I reflect on how I have grown both personally and scientifically since I interviewed with Leigh as a wide-eyed undergraduate student in fall 2015, I am filled with gratitude for the opportunities for learning and growth that Leigh, these whales, and many mentors and collaborators have offered over the years. As is often the case in science, the more questions you ask, the more questions you end up with. We are already dreaming up future studies to further understand the ecology, health, and resilience of this blue whale population. I can only imagine what we might learn in another decade.

Figure 5. A blue whale mother and calf pair come up for air in the South Taranaki Bight. Photo by Dawn Barlow.

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Barlow DR, Bernard KS, Escobar-Flores P, Palacios DM, Torres LG (2020) Links in the trophic chain: Modeling functional relationships between in situ oceanography, krill, and blue whale distribution under different oceanographic regimes. Mar Ecol Prog Ser 642:207–225.

Barlow DR, Estrada Jorge M, Klinck H, Torres LG (2022a) Shaken, not stirred: blue whales show no acoustic response to earthquake events. R Soc Open Sci 9:220242.

Barlow DR, Klinck H, Ponirakis D, Branch TA, Torres LG (2023) Environmental conditions and marine heatwaves influence blue whale foraging and reproductive effort. Ecol Evol 13:e9770.

Barlow DR, Klinck H, Ponirakis D, Garvey C, Torres LG (2021) Temporal and spatial lags between wind, coastal upwelling, and blue whale occurrence. Sci Rep 11:1–10.

Barlow DR, Klinck H, Ponirakis D, Holt Colberg M, Torres LG (2022b) Temporal occurrence of three blue whale populations in New Zealand waters from passive acoustic monitoring. J Mammal.

Barlow DR, Pepper AL, Torres LG (2019) Skin deep: An assessment of New Zealand blue whale skin condition. Front Mar Sci 6:757.

Barlow DR, Torres LG (2021) Planning ahead: Dynamic models forecast blue whale distribution with applications for spatial management. J Appl Ecol 58:2493–2504.

Barlow DR, Torres LG, Hodge KB, Steel D, Baker CS, Chandler TE, Bott N, Constantine R, Double MC, Gill P, Glasgow D, Hamner RM, Lilley C, Ogle M, Olson PA, Peters C, Stockin KA, Tessaglia-hymes CT, Klinck H (2018) Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger Species Res 36:27–40.

Torres LG (2013) Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zeal J Mar Freshw Res 47:235–248.

Torres LG, Barlow DR, Chandler TE, Burnett JD (2020) Insight into the kinematics of blue whale surface foraging through drone observations and prey data. PeerJ 8:e8906.

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

By Rachel Kaplan, PhD student, OSU College of Earth, Ocean and Atmospheric Sciences and Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab 

Ocean ecosystems are complex and dynamic, shaped by the interconnected physical and biogeochemical processes that operate across a variety of timescales. A trip on the “ocean conveyer belt”, which transports water from the North Atlantic across the global ocean and back in a process called thermohaline circulation, takes about a thousand years to complete. Phytoplankton blooms, which cycle nutrients through the surface ocean and feed marine animals, often occur at the crucial, food-poor moment of spring, and last for weeks or months. The entanglement of a whale in fishing gear, a major anthropogenic threat to ocean life that drives the GEMM Lab’s Project OPAL, can happen in seconds.

Compounding this complexity, even the timescales that research has clarified are changing. Many processes in the ocean are shifting – and often accelerating – due to global climate change. Images of melting sea ice, calving glaciers, and coastal erosion all exemplify our natural world’s rapid reorganization, and even discrete events can have dramatic repercussions and leave their mark for years. For example, a marine heatwave that occurred in 2014-2015 raised temperatures up to 2.5° C warmer than usual, redistributed species northward along the United States’ West Coast, spurred harmful algal blooms, and shut down fisheries. The toxic blooms also caused marine mammal strandings, domoic acid poisoning in California sea lions, and seabird mass death events (McCabe et al., 2016).

Figure 1. Figures like this Stommel diagram reveal the broad temporal and spatial scales over which ocean phenomena occur. Source: Sloyan et al., 2019

As humans seek to manage ocean ecosystems and mitigate the effects of climate change, our political processes have their own time scales, interconnected cycles, and stochasticity, just like the ocean. At the federal level in the United States, the legislative process takes place over months to decades, sometimes punctuated by relatively quicker actions enacted through Executive Orders. In addition, just as plankton have their turnover times, so do governmental branches. Both the legislative branch and the executive branch change frequently, with new members of Congress coming in every two years, and the president and administration changing every four or eight years. Turnover in both of these branches may constitute a total regime shift, with new members seeking to redirect science policy efforts.

The friction between oceanic and political timescales has historically made crafting effective ocean conservation policy difficult. In recent years, the policy approach of “adaptive management” has sought to respond to the challenges at the tricky intersection of politics, climate change, and ocean ecosystems. The U.S. Department of the Interior’s Technical Guide to Adaptive Management highlights its capacity to deal with the uncertainty inherent to changing ecosystems, and its ability to accommodate progress made through research: “Adaptive management [is a decision process that] promotes flexible decision making that can be adjusted in the face of uncertainties as outcomes from management actions and other events become better understood. Careful monitoring of these outcomes both advances scientific understanding and helps adjust policies or operations as part of an iterative learning process” (Williams et al, 2009).

Over the last several years, adaptive management policy approaches have been key as resource managers along the West Coast have responded to the problem of whale entanglement in fishing gear. When the 2014-2015 marine heatwave event caused anomalously low krill abundance in the central California Current region, humpback whales used a tactic called “prey-switching”, and fed on inshore anchovy schools rather than offshore krill patches. The resulting habitat compression fueled an increase in humpback whale entanglement events in Dungeness crab fishing gear (Santora et al, 2020). 

This sudden uptick in whale entanglements necessitated strategic management responses along the West Coast. In 2017, the California Dungeness Crab Fishing Gear Working Group developed the Risk Assessment and Mitigation Program (RAMP) to analyze real-time whale distribution and ocean condition data during the fishing season, and provide contemporaneous assessments of entanglement risk to the state’s Department of Fish and Wildlife. The Oregon Whale Entanglement Working Group (OWEWG) formed in 2017, tasked with developing options to reduce risk. Oregon Department of Fish and Wildlife (ODFW) has guided whale entanglement reduction efforts by identifying four areas of ongoing work: accountability, risk reduction, best management practices, and research – with regular, scheduled reviews of the regulations and opportunities to update and adjust them.

Figure 2. Entanglement in fishing gear can occur in seconds and may negatively impact whales for years. Source Scott Benson/NOAA

The need for research to support the best possible policy is where the GEMM Lab comes in. ODFW has established partnerships with Oregon State University and Oregon Sea Grant in order to improve understanding of whale distributions along the coast that can inform management efforts. Being involved in this cooperative “iterative learning process” is exactly why I’m so glad to be part of Project OPAL. Initial results from this work have already shaped ODFW’s regulations, and the framework of adaptive management and assessment means that regulations can continue being updated as we learn more through our research.

Ecosystem management will always be complex, just like ecosystems themselves. Today, the pace at which the climate is changing causes many people concern and even despair (Bryndum-Buchholz, 2022). Building adaptive approaches into marine policymaking, like the ones in use off the West Coast, introduces a new timescale into the U.S. policy cycle – one more in line with the rapid changes that are occurring within our dynamic ocean.



Williams, B. L., Szaro, R. C., and Shapiro, C. D. 2009. Adaptive management: the U.S. Department of the Interior Technical Guide. Adaptive Management Working Group, v pp.

Bryndum-Buchholz, A. (2022). Keeping up hope as an early career climate-impact scientist. ICES Journal of Marine Science, 79(9), 2345–2350. https://doi.org/10.1093/icesjms/fsac180

McCabe, R. M., Hickey, B. M., Kudela, R. M., Lefebvre, K. A., Adams, N. G., Bill, B. D., Gulland, F. M., Thomson, R. E., Cochlan, W. P., & Trainer, V. L. (2016). An unprecedented coastwide toxic algal bloom linked to anomalous ocean conditions. Geophys Res Lett, 43(19), 10366–10376. https://doi.org/10.1002/2016GL070023

Santora, J. A., Sydeman, W. J., Schroeder, I. D., Wells, B. K., & Field, J. C. (2011). Mesoscale structure and oceanographic determinants of krill hotspots in the California Current: Implications for trophic transfer and conservation. Progress in Oceanography, 91(4), 397–409. https://doi.org/10.1016/j.pocean.2011.04.002

Sloyan, B. M., Wilkin, J., Hill, K. L., Chidichimo, M. P., Cronin, M. F., Johannessen, J. A., Karstensen, J., Krug, M., Lee, T., Oka, E., Palmer, M. D., Rabe, B., Speich, S., von Schuckmann, K., Weller, R. A., & Yu, W. (2019). Evolving the Physical Global Ocean Observing System for Research and Application Services Through International Coordination. Frontiers in Marine Science, 6, 449. https://doi.org/10.3389/fmars.2019.00449

Decisions, decisions: New GEMM Lab publication reveals trade-offs in prey quantity and quality in gray whale foraging

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

Obtaining enough food is crucial for predators to ensure adequate energy gain for maintenance of vital functions and support for energetically costly life history events (e.g., reproduction). Foraging involves decisions at every step of the process, including prey selection, capture, and consumption, all of which should be as efficient as possible. Making poor foraging decisions can have long-term repercussions on reproductive success and population dynamics (Harris et al. 2007, 2008, Grémillet et al. 2008), and for marine predators that rely on prey that is spatially and temporally dynamic and notoriously patchy (Hyrenbach et al. 2000), these decisions can be especially challenging. Prey abundance and density are frequently used as predictors of marine predator distribution, movement, and foraging effort, with predators often selecting highly abundant or dense prey patches (e.g., Goldbogen et al. 2011, Torres et al. 2020). However, there is increased recognition that prey quality is also an important factor to consider when assessing a predator’s ecology and habitat use (Spitz et al. 2012), and marine predators do show a preference for higher quality prey items (e.g., Haug et al. 2002, Cade et al. 2022). Moreover, negative impacts of low-quality prey on the health and breeding success of some marine mammals (Rosen & Trites 2000, Trites & Donnelly 2003) have been documented. Therefore, examining multiple prey metrics, such as prey quantity and quality, in predator ecology studies is critical.

Figure 1. Site map of the Port Orford TOPAZ/JASPER integrated projects. Blue squares represent the location of the 12 sampling stations within the 2 study sites (site boundaries demarcated with black lines). Brown dot represents the cliff-top observation site where theodolite tracking occurred.

Our integrated TOPAZ/JASPER projects in Port Orford do just this! We collect both prey quantity and quality data from a tandem research kayak, while we track Pacific Coast Feeding Group (PCFG) gray whales from shore. The prey and whale sampling overlap spatially (and often temporally within the same day). This kind of concurrent predator-prey sampling at similar scales is often logistically challenging to achieve, yet because PCFG gray whales have an affinity for nearshore, coastal habitats, it is something we have been able to achieve in Port Orford. Since 2016, a field team comprised of graduate, undergraduate, and high school students has collected data during the month of August to investigate gray whale foraging decisions relative to prey. Every day, a kayak team collects GoPro videos (to assess relative prey abundance; AKA: quantity) and zooplankton samples using a tow net (to assess prey community composition; AKA: quality through caloric content of different species) (Figure 1). At the same time, a cliff team surveys for gray whales from shore and tracks them using a theodolite, which provides us with tracklines of individual whales; We categorize each location of a whale into three broad behavior states (feeding, searching, transiting) based on movement patterns. Over the years, the various students who have participated in the TOPAZ/JASPER projects have written many blog posts, which I encourage you to read here (particularly to get more detailed information about the field methods). 

Figure 2. An example daily layer of relative prey abundance (increasing color darkness corresponds with increasing abundance) in one study site with a whale theodolite trackline recorded on the same day overlaid and color-coded by behavioral state.

Several years of data are needed to conduct a robust analysis for our ecological questions about prey choice, but after seven years, we finally had the data and I am excited to share the results, which are due to the many years of hard work from many students! Our recent paper in Marine Ecology Progress Series aimed to determine whether PCFG gray whale foraging decisions are driven by prey quantity (abundance) or quality (caloric content of species) at a scale of 20 m (which is slightly less than 2 adult gray whale body lengths). In this study, we built upon results from my previous Master’s publication, which revealed that there are significant differences in the caloric content between the six common nearshore zooplankton prey species that PCFG gray whales feed on (Hildebrand et al. 2021). Therefore, in this study we addressed the hypothesis that individual whales will select areas where the prey community is dominated by the mysid shrimp Neomysis rayii, since it is significantly higher in caloric content than the other two prey species we identified, Holmesimysis sculpta (a medium quality mysid shrimp species) and Atylus tridens (a low quality amphipod species) (Hildebrand et al. 2021). We used spatial statistics and model to make daily maps of prey abundance and quality that we compared to our whale tracks and behavior from the same day. Please read our paper for the details on our novel methods that produced a dizzying amount of prey layers, which allowed us to tease apart whether gray whales target prey quantity, quality, or a mixture of both when they forage. 

Figure 3. Figure shows the probability of gray whale foraging relative to prey abundance (color-coded by prey species). Dark grey vertical line represents the mean threshold for the H. sculpta curves (12.0); light grey vertical lines: minimum (7.2) and maximum (15.3) thresholds for the H. sculpta curves. Inflection points could not be calculated for the N. rayii curves

So, what did we find? The models proved our hypothesis wrong: foraging probability was significantly correlated with the quantity and quality of the mysid H. sculpta, which has significantly lower calories than N. rayii. This result puzzled us, until we started looking at the overall quantity of these two prey types in the study area and realized that the amount of calorically-rich N. rayii never reached a threshold where it was beneficial for gray whales to forage. But, there was a lot of H. sculpta, which likely made for an energetic gain for the whales despite not being as calorically rich as N. rayii. We determined a threshold of H. sculpta relative abundance that is required to initiate the gray whale foraging behavior, and the abundance of N. rayii never came close to this level (Figure 3). Despite not having the highest quality, H. sculpta did have the highest abundance and showed a significant positive relationship with foraging behavior, unlike the other prey items. Interestingly, whales never selected areas dominated by the low-calorie species A. tridens. These results demonstrate trade-off choices by whales for this abundant, medium-quality prey.

To our knowledge, individual baleen whale foraging decisions relative to available prey quantity and quality have not been addressed previously at this very fine-scale. Interestingly, this trade-off between prey quantity and quality has also been detected in humpback whales foraging in Antarctica at depths deeper than where the densest krill patches occur; while the whales are exploiting less dense krill patches, these krill composed of larger, gravid, higher-quality krill (Cade et al. 2022). While it is unclear how baleen whales differentiate between prey species or reproductive stages, several mechanisms have been suggested, including visual and auditory identification (Torres 2017). We assume here that gray whales, and other baleen whale species, can differentiate between prey species. Thus, our results showcase the importance of knowing the quality (such as caloric content) of prey items available to predators to understand their foraging ecology (Spitz et al. 2012). 


Cade DE, Kahane-Rapport SR, Wallis B, Goldbogen JA, Friedlaender AS (2022) Evidence for size-selective pre- dation by Antarctic humpback whales. Front Mar Sci 9:747788

Goldbogen JA, Calambokidis J, Oleson E, Potvin J, Pyenson ND, Schorr G, Shadwick RE (2011) Mechanics, hydrody- namics and energetics of blue whale lunge feeding: effi- ciency dependence on krill density. J Exp Biol 214:131−146

Grémillet D, Pichegru L, Kuntz G, Woakes AG, Wilkinson S, Crawford RJM, Ryan PG (2008) A junk-food hypothesis for gannets feeding on fishery waste. Proc R Soc B 275: 1149−1156

Harris MP, Beare D, Toresen R, Nøttestad L, and others (2007) A major increase in snake pipefish (Entelurus aequoreus) in northern European seas since 2003: poten- tial implications for seabird breeding success. Mar Biol 151:973−983

Harris MP, Newell M, Daunt F, Speakman JR, Wanless S (2008) Snake pipefish Entelurus aequoreus are poor food for seabirds. Ibis 150:413−415

Haug T, Lindstrøm U, Nilssen KT (2002) Variations in minke whale (Balaenoptera acutorostrata) diet and body condi- tion in response to ecosystem changes in the Barents Sea. Sarsia 87:409−422

Hildebrand L, Bernard KS, Torres LG (2021) Do gray whales count calories? Comparing energetic values of gray whale prey across two different feeding grounds in the eastern North Pacific. Front Mar Sci 8:1008

Hyrenbach KD, Forney KA, Dayton PK (2000) Marine pro- tected areas and ocean basin management. Aquat Con- serv 10:437−458

Rosen DAS, Trites AW (2000) Pollock and the decline of Steller sea lions: testing the junk-food hypothesis. Can J Zool 78:1243−1250

Spitz J, Trites AW, 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:e50096

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Torres LG (2017) A sense of scale: foraging cetaceans’ use of scale-dependent multimodal sensory systems. Mar Mamm Sci 33:1170−1193

Trites AW, Donnelly CP (2003) The decline of Steller sea lions Eumetopias jubatus in Alaska: a review of the nutri- tional stress hypothesis. Mammal Rev 33:3−28

A Hundred and One Data Visualizations: What We Can Infer about Gray Whale Health Using Public Data

By Braden Adam Vigil, Oregon State University Undergraduate, GEMM Lab NSF REU Intern


My name is Braden Vigil, and I am enjoying this summer with the company of Lisa Hildebrand and Dr. Leigh Torres as a National Science Foundation (NSF) Research Experience for Undergraduates (REU) intern. By slicing off a manageable chunk of the GRANITE project to focus on, I’ve had the chance to explore my passion for data visualization. My excitement for biological research was instilled in me by an impactful high school biology teacher (thank you Mr. Villalobos!) and was narrowed to marine biology research after a chance visit to Oregon State University’s Hatfield Marine Science Center. I’ve come all the way from Southern New Mexico to explore this passion of mine, and the REU program has been one of my first chances to get my feet wet. My advice for any students debating taking big leaps for the sake of passion is to do it – it’s scary, but I’d say there’s nothing better than living out what you want to do (and hopefully getting paid for it!). For this project, the GEMM Lab has saved me the trouble of collecting data – this summer, I’m all action. 

Where Gray Whales Are and Why It Matters

Just as you might find yourself at a grocery store to buy food or at a coffee shop catching up with an old friend, so too do whales have places to go and reasons for being there. Research concerning gray whale ecology – understanding the who, what, when, where, whys – should then have a lot to do with the “where?” and “why?” That’s what my project is about: investigating where the gray whales off the Oregon coast are, and what features of the environment are related to their presence and other aspects of the population. After all, distribution is considered the foundational unit for the biogeographical understanding of a population’s location and its interactions with other species. An example of an environmental driver may be phytoplankton and – subsequently – zooplankton abundance. It’s been shown that bottom-up trophic cascades based on primary productivity directly influence predator and prey populations in both terrestrial and marine ecosystems (Sinclair and Krebs 2002; Benoit-Bird and McManus 2012). This driver specifically could then inform something as significant as population abundance of a predator, though that’s out of the scope of my project. Instead, I’m studying how these environmental drivers, specifically sea water temperature, affects the variation of the thyroid hormone (tri-iodothyronine, T3) in gray whales, which the GEMM Lab quantifies from fecal samples that they non-invasively and opportunistically collect. In terrestrial mammals, T3 is believed to be associated with thermoregulation, yet it is unclear if T3 has the same function in baleen whales who use blubber insulation to thermoregulate. To estimate blubber insulation, we use a proxy, called body area index (BAI) collected via drone footage (Burnett et al. 2018), which you can read more about in Clara’s blog. Insights into variations in T3 hormone levels as related to changes in the environment may allow researchers to better understand thermoregulatory challenges whales face in warming oceans.

This Sounds Like a Lot of Data About the Environment, Where’s it Coming From?

Not only has the GEMM Lab relieved me of the hassle that data collection and fieldwork can be, so too has the Ocean Observatories Initiative (OOI). Starting in 2014, the OOI has set up several buoys off the U.S. West Coast, each equipped with numerous sensors and data-collecting devices. These have been extracting data from the nearby environment since then, including aspects such as dissolved oxygen, pH, and most important to this study, sea temperature. These buoys run deep too! Some devices reach as low as 25 m, which is where we often expect to see whales foraging during surveys. For our interest, there is one specific buoy that is within the GRANITE project’s survey region, the Oregon Inshore Surface Mooring.

Figure 1. Locations of OOI buoys. Blue dots represent buoys, while the yellow dot represents our buoy of interest, the Oregon Inshore Surface Mooring. 


The OOI has published, and continues to publish, an unbelievable amount of data. There are many things that would be interesting to investigate, but until we know how much we can bite off versus how much we can chew, we’ve narrowed it down to a few hypotheses we’re currently investigating. 

Table 1. Hypotheses and Expected Results.

A Hundred and One Data Visualizations

As fun as I find testing correlations between variables and creating satisfying looking plots, I must admit that I’m not even halfway into this project and I’ve made a LOT of plots. Plots can be an easy way to understand big datasets and observations. Since not all of the data-collecting devices on the OOI data are continuously running, I first needed to get an idea of how much data we have to work with, and how much of that data overlaps in time with our annual gray whale survey period (June 1 – October 15). Some of these preliminary plots look like Figure 2. In addition, these plots grant us an idea of how variable sea surface temperatures have been in these past few years. Marine heatwaves have occurred recently in the Pacific Ocean and off the U.S. West Coast, and it is important to know if their effects continue to linger to the present. Other, unexplained peaks might also be worth investigating. 

Figure 2. Preliminary plot comparing sea surface temperature data over time, from around June 2016 to December 2021. Straight lines between December to June each year indicates no data, as we have removed these periods from our analysis. 

The goal here is to eventually compare the variables of sea temperature to the T3 hormone levels in gray whales foraging off the Oregon coast. Before this step, it is important to decide what depth of temperature readings are most appropriate to assess. I’ve made several correlation plots of sea  temperature between varying depths of 1 m, 7 m, and 25 m. One such plot is included below (Figure 3). This plot shows variation of temperature between different depths. If there is strong variation between the depths of 1 m and 25 m, then the water column may be well stratified, meaning that gray whale response to environmental temperature may be distinct between these distances, possibly even between 1 m and 7 m. 

Figure 3. Sea surface temperature at 1 m versus 25 m in degrees Celsius, with points color coded by year. 


As previously described, this study plays part into the larger GRANITE project with the goal to understand and make predictions about the ecology and physiology of the gray whale population off of the U.S. West Coast. This study will investigate the significance of sea temperature on aspects of whale health – so far including BAI and T3 hormone level. I will be pursuing a stronger grasp on the variation of these relationships through ongoing analysis. My results should be used to clarify nodes and the correlation between them in the web of dynamics encircling the population. This project has given me great insight into how raw data can be turned into meaningful understandings and subsequent impacts. The public OOI data is a scattershot of many different measurements using many different devices constantly. The answers/solutions to the conservation of species threatened by the Anthropocene are out there, all that’s required is that we harness them. 


Benoit-Bird, K. J., & McManus, M. A. (2012). Bottom-up regulation of a pelagic community through spatial aggregations. Biology Letters8(5), 813–816. https://doi.org/10.1098/rsbl.2012.0232

Burnett, J. D., & Wing, M. G. (2018). A low-cost near-infrared digital camera for fire detection and monitoring. International Journal of Remote Sensing39(3), 741–753. https://doi.org/10.1080/01431161.2017.1385109

Sinclair, A. R. E., & Krebs, C. J. (2002). Complex numerical responses to top–down and bottom–up processes in vertebrate populations. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences357(1425), 1221–1231.https://doi.org/10.1098/rstb.2002.1123.