Rock-solid GRANITE: Scaling the disturbance response of individual whales up to population level impacts

Since early May, much of the GEMM Lab has been consumed by the GRANITE project, which stands for Gray whale Response to Ambient Noise Informed by Technology and Ecology. Two weeks ago, PhD student Clara Bird discussed our field work preparations, and since May 20th we have conducted five successful days of field work (and one unsuccessful day due to fog). If you are now expecting a blog about the data we have collected so far and whales we encountered, I am sorry to disappoint you. Rather, I want to take a big step back and provide the context of the GRANITE project as a whole, explain why this project and data collection is so important, and discuss what it is that we hope to achieve with our ever-growing, multidisciplinary dataset and team.

We use the Pacific Coast Feeding Group (PCFG) of gray whales that forage off the Oregon coast as our study system to better understand the ecological and physiological response of baleen whales to multiple stressors. Our field methodology includes replicate physiological and ecological sampling of this accessible baleen whale population with synoptic measurement of multiple types of stressors. We collect fecal samples for hormone analysis, conduct drone overflights of whales to collect body condition and behavioral data, record the ambient soundscape through deployment of two hydrophones, and conduct whale photo-identification to link all data streams to each individual whale of known sex, estimated age, and reproductive status. We resample these data from multiple individuals within and between summer foraging seasons, while exposed to different potential stressors occurring at different intensities and temporal periods and durations. The hydrophones are strategically placed with one in a heavily boat-trafficked (and therefore noisy) area close to the Port of Newport, while the second is located in a relatively calm (and therefore quieter) spot near the Otter Rock Marine Reserve (Fig. 1). These hydrophones provide us with information about both natural (e.g. killer whales, wind, waves) and anthropogenic (e.g. boat traffic, seismic survey, marine construction associated with PacWave wave energy facility development) noise that may affect gray whales. During sightings with whales, we also drop GoPro cameras and sample for prey to better understand the habitats where whales forage and what they might be consuming.

Figure 1. Map of GRANITE study area from Seal Rock to Lincoln City with gray whale sightings (yellow circles) and and fecal samples collected (red triangles) from the 2020 field season. Green stars represent the two hydrophone locations. Source: L. Torres.

GEMM Lab PI Dr. Leigh Torres initiated this research project in 2015 and established partnerships with acoustician Dr. Joe Haxel and (then) PhD student Dr. Leila Lemos. Since then, the team working on this project has grown considerably to provide expertise in the various disciplines that the project integrates. Leigh is currently joined at the GRANITE helm by 4 co-PIs: Dr. Haxel, endocrinologist Dr. Kathleen Hunt, biological statistician Dr. Leslie New, and physiologist Dr. Loren Buck. Drs. Alejandro Fernandez Ajo, KC Bierlich and Enrico Pirotta are postdoctoral scholars who are working on the endocrinology, photogrammetry, and biostatistical modelling components, respectively. Finally, Clara and myself are partially funded through this project for our PhD research, with Clara focusing on the links between behavior, body condition, individualization, and habitat, while I am tackling questions about the recruitment and site fidelity of the PCFG (more about these topics below). 

Faculty Research Assistant Todd Chandler supervises PhD student Clara Bird during her maiden drone flight over a whale. Source: L. Torres.

The ultimate goal of this project is to use the PCFG as a case study to quantify baleen whale physiological response to different stressors and model the subsequent impacts on the population by implementing our long-term, replicate dataset into a framework called Population consequences of disturbance (PCoD; Fig. 2). PCoD is built upon the underlying concept that changes in behavior and/or physiology caused by disturbance (i.e. noise) affect the fitness of individuals by impacting their health and vital rates, such as survival, reproductive success, and growth rate (Pirotta et al. 2018). These impacts at the individual level may (or may not) affect the population as a whole, depending on what proportion of individuals in the population are affected by the disturbance and the intensity of the disturbance effect on each individual. The PCoD framework requires quantification of four stages: a) the physiological and/or behavioral changes that occur as a result of exposure to a stressor (i.e. noise), b) the acute effects of these physiological and/or behavioral responses on individual vital rates, and their chronic effects via individual health, c) the way in which changes in health may affect the vital rates of individuals, and d) how changes in individual vital rates may affect population dynamics (Fig. 2; Pirotta et al. 2018). While four stages may not sound like a lot, the amount and longevity of data needed to quantify each stage is immense. 

Figure 2. Conceptual framework of the population consequences of disturbance (PCoD). Letters (A-D) represent the four stages that require quantification in order for PCoD to be implemented. Each colored box represents external (ecological drivers, stressors) and internal (physiology, health, vital rates, behavior) factors that can change over time that are measured for each individual whale (dashed grey boundary line). The effects are then integrated across all individuals in the population to project their effects on the population’s dynamics. Figure and caption adapted from Pirotta et al. 2018.

The ability to detect a change in behavior or physiology often requires an understanding of what is “normal” for an individual, which we commonly refer to as a baseline. The best way to establish a baseline is to collect comprehensive data over a long time period. With our data collection efforts since 2015 of fecal samples, drone flights and photo identification, we have established useful baselines of behavioral and physiological data for PCFG gray whales. These baselines are particularly impressive since it is typically difficult to collect repeated measurements of hormones and body condition from the same individual baleen whale across multiple years. These repeated measurements are important because, like all mammals, hormones and body condition vary across life history phases (i.e., with pregnancy, injury, or age class) and across time (i.e., good or bad foraging conditions). To achieve these repeated measurements, GRANITE exploits the high degree of intra- and inter-annual site fidelity of the PCFG, their accessibility for study due to their affinity for nearshore habitat use, and the long-term sighting history of many whales that provides sex and approximate age information. Our work to-date has already established a few important baselines. We now know that the body condition of PCFG gray whales increases throughout a foraging season and can fluctuate considerably between years (Soledade Lemos et al. 2020). Furthermore, there are significant differences in body condition by reproductive state, with calves and pregnant females displaying higher body conditions (Soledade Lemos et al. 2020). Our dataset has also allowed us to validate and quantify fecal steroid and thyroid hormone metabolite concentrations, providing us with putative thresholds to identify a stressed vs. not stressed whale based on its hormone levels (Lemos et al. 2020).

PhD student Lisa Hildebrand and GRANITE co-PI Dr. Kathleen Hunt collecting a fecal sample. Source: L. Torres.

We continue to collect data to improve our understanding of baseline PCFG physiology and behavior, and to detect changes in their behavior and physiology due to disturbance events. All these data will be incorporated into a PCoD framework to scale from individual to population level understanding of impacts. However, more data is not the only thing we need to quantify each of the PCoD stages. The implementation of the PCoD framework also depends on understanding several aspects of the PCFG’s population dynamics. Specifically, we need to know whether recruitment to the PCFG population occurs internally (calves born from “PCFG mothers” return to the PCFG) or externally (immigrants from the larger Eastern North Pacific gray whale population joining the PCFG as adults). The degree of internal or external recruitment to the PCFG population should be included in the PCoD model as a parameter, as it will influence how much individual level disturbance effects impact the overall health and viability of the population. Furthermore, knowing residency times and home ranges of whales within the PCFG is essential to understand exposure durations to disturbance events. 

To assess both recruitment and residency patterns of the PCFG, I am undertaking a large photo-identification effort, which includes compiling sightings and photo data across many years, regions, and collaborators. Through this effort we aim to identify calves and their return rate to the population, the rate of new adult recruits to the population, and the spatial residency of individuals in our study system. Although photo-id is a basic, commonplace method in marine mammal science, its role is critical to tracking individuals over time to understand population dynamics (in a non-invasive manner, no less). A large portion of my PhD research will focus on the tedious yet rewarding task of photo-id data management and matching in order to address these pressing knowledge gaps on PCFG population dynamics needed to implement the PCoD model that is an ultimate goal of GRANITE. I am just beginning this journey and have already pinpointed many analytical and logistical hurdles that I need to overcome. I do not anticipate an easy path to addressing these questions, but I am extremely eager to dig into the data, reveal the patterns, and integrate the findings into our rock-solid GRANITE project.  

Funding for the GRANITE project comes from the Office of Naval Research, the Department of Energy, Oregon Sea Grant, the NOAA/NMFS Ocean Acoustics Program, and the OSU Marine Mammal Institute.


Lemos, L.S., Olsen, A., Smith, A., Chandler, T.E., Larson, S., Hunt, K., and L.G. Torres. 2020. Assessment of fecal steroid and thyroid hormone metabolites in eastern North Pacific gray whales. Conservation Physiology 8:coaa110.

Pirotta, E., Booth, C.G., Costa, D.P., Fleishman, E., Kraus, S.D., Lusseau, D., Moretti, D., New, L.F., Schick, R.S., Schwarz, L.K., Simmons, S.E., Thomas, L., Tyack, P.L., Weise, M.J., Wells, R.S., and J. Harwood. 2018. Understanding the population consequences of disturbance. Ecology and Evolution 8(19):9934-9946.

Soledade Lemos, L., Burnett, J.D., Chandler, T.E., Sumich, J.L., and L.G. Torres. 2020. Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11(4):e03094.

The learning curve never stops as the GRANITE project begins its seventh field season

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

When I thought about what doing fieldwork would be like, before having done it myself, I imagined that it would be a challenging, but rewarding and fun experience (which it is). However, I underestimated both ends of the spectrum. I simultaneously did not expect just how hard it would be and could not imagine the thrill of working so close to whales in a beautiful place. One part that I really did not consider was the pre-season phase. Before we actually get out on the boats, we spend months preparing for the work. This prep work involves buying gear, revising and developing protocols, hiring new people, equipment maintenance and testing, and training new skills. Regardless of how many successful seasons came before a project, there are always new tasks and challenges in the preparation phase.

For example, as the GEMM Lab GRANITE project team geared up for its seventh field season, we had a few new components to prepare for. Just to remind you, the GRANITE (Gray whale Response to Ambient Noise Informed by Technology and Ecology) project’s field season typically takes place from June to mid-October of each year. Throughout this time period the field team goes out on a small RHIB (rigid hull inflatable boat), whenever the weather is good enough, to collect photo-ID data, fecal samples, and drone imagery of the Pacific Coast Feeding Group (PCFG) gray whales foraging near Newport, OR, USA. We use the data to assess the health, ecology and population dynamics of these whales, with our ultimate goal being to understand the effect of ambient noise on the population. As previous blogs have described, a typical field day involves long hours on the water looking for whales and collecting data. This year, one of our exciting new updates is that we are going out on two boats for the first part of the field season and starting our season 10 days early (our first day was May 20th). These updates are happening because a National Science Foundation funded seismic survey is being conducted within our study area starting in June. The aim of this survey is to assess geophysical structures but provides us with an opportunity to assess the effect of seismic noise on our study group by collecting data before, during, and after the survey. So, we started our season early in order to capture the “before seismic survey” data and we are using a two-boat approach to maximize our data collection ability.

While this is a cool opportunistic project, implementing the two-boat approach came with a new set of challenges. We had to find a second boat to use, buy a new set of gear for the second boat, figure out the best way to set up our gear on a boat we had not used before, and update our data processing protocols to include data collected from two boats on the same day. Using two boats also means that everyone on the core field team works every day. This core team includes Leigh (lab director/fearless leader), Todd (research assistant), Lisa (PhD student), Ale (new post-doc), and me (Clara, PhD student). Leigh and Todd are our experts in boat driving and working with whales, Todd is our experienced drone pilot, I am our newly certified drone pilot, and Lisa, Ale, and myself are boat drivers. Something I am particularly excited about this season is that Lisa, Ale, and I all have at least one field season under our belts, which means that we get to become more involved in the process. We are learning how to trailer and drive the boats, fly the drones, and handling more of the post-field work data processing. We are becoming more involved in every step of a field day from start to finish, and while it means taking on more responsibility, it feels really exciting. Throughout most of graduate school, we grow as researchers as we develop our analytical and writing skills. But it’s just as valuable to build our skillset for field work. The ocean conditions were not ideal on the first day of the field season, so we spent our first day practicing our field skills.

For our “dry run” of a field day, we went through the process of a typical day, which mostly involved a lot of learning from Leigh and Todd. Lisa practiced her trailering and launching of the boat (figure 1), Ale and Lisa practiced driving the boat, and I practiced flying the drone (figure 2). Even though we never left the bay or saw any whales, I thoroughly enjoyed our dry run. It was useful to run through our routine, without rushing, to get all the kinks out, and it also felt wonderful to be learning in a supportive environment. Practicing new skills is stressful to say the least, especially when there is expensive equipment involved, and no one wants to mess up when they’re being watched. But our group was full of support and appreciation for the challenges of learning. We cheered for successful boat launchings and dockings, and drone landings. I left that day feeling good about practicing and improving my drone piloting skills, full of gratitude for our team and excited for the season ahead.

Figure 1. Lisa (driving the truck) launching the boat.
Figure 2. Clara (seated, wearing a black jacket) landing the drone in Ale’s hands.

All the diligent prep work paid off on Saturday with a great first day (figure 3). We conducted five GoPro drops (figure 4), collected seven fecal samples from four different whales (figure 5), and flew four drone flights over three individuals including our star from last season, Sole. Combined, we collected two trifectas (photo-ID images, fecal samples, and drone footage)! Our goal is to get as many trifectas as possible because we use them to study the relationship between the drone data (body condition and behavior) and the fecal sample data (hormones). We were all exhausted after 10 hours on the water, but we were all very excited to kick-start our field season with a great day.

Figure 3. Lisa on the bow pulpit during our first sighting of the day.
Figure 4. Lisa doing a GoPro drop, she’s lowering the GoPro into the water using the line in her hands.
Figure 5. Clara and Ale collecting a fecal sample.

On Sunday, just one boat went out to collect more data from Sole after a rainy morning and I successfully flew over her from launching to landing! We have a long season ahead, but I am excited to learn and see what data we collect. Stay tuned for more updates from team GRANITE as our season progresses!

Why did I start sketchnoting?

By Solène Derville, Postdoc, OSU Department of Fisheries, Wildlife, and Conservation Science, Geospatial Ecology of Marine Megafauna Lab

Sketchnoting, also known as « visual notetaking » is a technique combining words with drawings, diagrams and typography to record ideas (Figure 1). This concept was invented by designer Michael Rohde in 2006 to combine tedious notetaking with doodling. He quickly discovered that adding drawings to his notes helped him concentrate and remember better. He would also be more likely to come back to his notes later on (something we must all admit is not so common). Similarly, after I followed a short online class by Magalie Le Gall (Sorbonne Université) I became convinced that sketchnoting shows  promise and can have a positive impact on my scientific work.

Figure 1 : What is sketchnoting ? By

Draw to remember more

The impact of sketchnoting on memory is not without scientific backing. Back in 1971, Allan Paivio, an American professor of psychology, developed the dual-coding theory. It posits that visual and verbal information are mentally processed in two distinctive systems and have additive effects on cognitive operations such as memory. Numerous experiments have empirically confirmed that dual coding (images + words) improve learning and memory. In addition, converting what you hear or see into visually interconnected drawings and words helps you synthesize content. Personalizing ideas into your own symbols and images also lays a strong basis for remembering. The implications of sketchnoting for educational purposes are therefore huge!

Draw to stay focused

I have only started sketchnoting recently but the impact this method had on my concentration immediately struck me. In the constant stream of information that we experience nowadays, I found that synthesizing ideas on paper using symbols and diagrams helped me stay focused on what I am presently reading or hearing, instead of letting my thoughts drift in a thousand different directions. Again, this outcome can have big implications in the classroom or at your desk. Using very basic lettering, bullets, frames and connectors (Figure 2), sketchnoting appears to be a good didactic tool.

Figure 2 : A few drawing tips by sketchnoter Carol Anne McGuire.

Draw to create and appeal

Figure 3 (source: ASIDE 2013)

Mike Rohde’s motto is « ideas, not art » because a lot of people have an immediate reaction of fear of failure when they are asked to draw something. He emphasizes that sketchnoting is not necessarily meant to be pretty, as it mostly serves a personal purpose. However, if you have an artistic fiber (even slightly!), sketchnoting becomes a great communication tool and can help you convey ideas in posters, slides, blogs, etc. Even very simple drawings are appealing and fun. You can create your own visual libraries from a few basic shapes (Figure 3). Anything can be drawn with a few simple lines! You can also use drawing libraries such as to find examples and eventually gain confidence… as you realize that the average people’s drawing skills are pretty low (the dolphin drawings on this website are worth a look)!

Now, the key to developing this new skill is clearly to practice! From now on, I have decided to record every one of our monthly GEMM lab meetings in a sketchnote to make sure I keep track of our great discussions. I will also definitely try to apply this approach when reading scientific literature, attending conferences, preparing drafts, teaching and so much more! And for a start, what could be better then to sketchnote the research project I currently working on (Figure 4)?

Figure 4 : My first attempt at sketchnoting! Illustration of the OPAL project that I am working on (credit : S. Derville).

References & resources:

Great intro to sketchnoting by Mike Rhode:

Training, tips, videos etc.:

Link to many ressources and websites:

Paivio, A (1971). Imagery and verbal processes. New York: Holt, Rinehart, and Winston.

Into the Krillscape: A Remote Expedition in Research and Mentorship

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

What are the most unexpected things you’ve done on Zoom in the last year? Since the pandemic dramatically changed all our lives in 2020, I think we’ve all been surprised by the diversity of things we’ve done remotely. I’ve baked bagels with a friend in Finland, done oceanography labs from my kitchen, had dance parties with people across the country, and conducted an award ceremony for my family’s Thanksgiving scavenger hunt – all on Zoom. Over the last several months, I’ve also mentored an Undergraduate Research, Scholarship, & the Arts (URSA) Engage student, named Amanda. Although we haven’t met in person yet, we’ve been connecting over Zoom since October. 

Amanda is an Ocean Sciences student working with me and Dr. Kim Bernard (CEOAS) to conduct a literature review about the two species of krill found off the coast of Oregon. Thysanoessa spinifera and Euphausia pacifica are an important food source for many of the animals that live off our coast — including blue, humpback, and fin whales. I am trying to learn how krill distributions shape those of humpback and blue whales as part of project OPAL, as well as which oceanographic factors drive krill abundances and distributions.

Thysanoessa spinifera (source: Scripps Institute of Oceanography). 

We’re also interested in T. spinifera and E. pacifica for the crucial roles they serve in ecosystems, beyond providing dinner for whales. Krill do many things that are beneficial to ecosystems and people, termed “ecosystem services.” These include facilitating carbon drawdown from the surface ocean to the deep, supporting lucrative fisheries species like salmon, flatfish, and rockfish, and feeding seabirds like auklets and shearwaters. We want to understand more fully the niche that T. spinifera and E. pacifica each fill off the coast of Oregon, which will help us anticipate how these important animals can be impacted by forces such as global climate change and marine management efforts.

Trying to understand the ecosystem services fulfilled by krill is inherently interdisciplinary, which means we have to learn a lot of new things, making this project a lot of fun. The questions Amanda and I have pursued together have ranged from intensely specific, to surprisingly broad. How many calories do blue whales need to eat in a day? How many krill do salmon need to eat? How big are krill fecal pellets, and how fast do they sink?

Trying to answer these questions has basically amounted to a heroic scouring of the internet’s krillscape by Amanda. She has hunted down papers dating back to the 1960s, pulled together findings from every corner of the world, and pursued what she refers to as “treasure troves” of data. In the process, she has also revealed the holes that exist in the literature, and given us new questions. This is the basis of the scientific process: understanding the current state of knowledge, identifying gaps in that knowledge, and developing the questions and methods needed to fill those gaps.

Euphausia pacifica (source: University of Irvine California, Peter J. Bryant).

Filling in knowledge gaps about T. spinifera and E. pacifica can help us better understand these animals, the ecosystems where they live, and the whales and other animals that depend on them for prey. It’s exciting to know that we will have the opportunity to help fill some of these gaps, as both Amanda and I continue this research over the course of our degrees.

Being able to engage in remote research and mentorship has been really rewarding, and it has shown me how far we’ve all come over the last year. Learning how to work together remotely has been crucial as we have adjusted to the funny new normal of the pandemic. As much as I miss working with people in person, I’ve learned that there’s a lot of great connection to be found even in remote collaboration – I’ve loved meeting Amanda’s pets on Zoom, learning about her career goals, and seeing her incredibly artistic representations of the carbon cycle held up to the camera.

Even though most of our conversations take place on Zoom from our homes, this research still feels plugged into a bigger community. Amanda and I also join Kim’s bigger Zooplankton Ecology Lab meetings, which include two other graduate students and eight undergraduate students, all of whom are working on zooplankton ecology questions that span from the Arctic to the Antarctic. Even though we’ve never met in person, a supportive and curious community has developed among all of us, which I know will persist when we can move back to in-person research and mentorship.

The right tool for the job: examining the links between animal behavior, morphology and habitat

Clara Bird, PhD Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

In order to understand a species’ distribution, spatial ecologists assess which habitat characteristics are most often associated with a species’ presence. Incorporating behavior data can improve this analysis by revealing the functional use of each habitat type, which can help scientists and managers assign relative value to different habitat types. For example, habitat used for foraging is often more important than habitat that a species just travels through. Further complexity is added when we consider that some species, such as gray whales, employ a variety of foraging tactics on a variety of prey types that are associated with different habitats. If individual foraging tactic specialization is present, different foraging habitats could be valuable to specific subgroups that use each tactic. Consequently, for a population that uses a variety of foraging tactics, it’s important to study the associations between tactics and habitat characteristics.

Lukoschek and McCormick’s (2001) study investigating the spatial distribution of a benthic fish species’ foraging behavior is a great example of combining data on behavior, habitat, and morphology.  They collected data on the diet composition of individual fish categorized into different size classes (small, medium, and large) and what foraging tactics were used in which reef zones and habitat types. The foraging tactics ranged from feeding in the water column to digging (at a range of depths) in the benthic substrate. The results showed that an interesting combination of fish behavior and morphology explained the observed diet composition and spatial distribution patterns. Small fish foraged in shallower water, on smaller prey, and primarily employed the water column and shallow digging tactics. In contrast, large fish foraged in deep water, on larger prey, and primarily fed by digging deeper into the seafloor (Figure 1). This pattern is explained by both morphology and behavior. Morphologically, the size of the feeding apparatus (mouth gape size) affects the size of the prey that a fish can feed on. The gape of the small fish is not large enough to eat the larger prey that large fish are able to consume. Behaviorally, predation risk also affects habitat selection and tactic use. Small fish are at higher risk of being predated on, so they remain in shallow areas where they are more protected from predators and they don’t dig as deep to forage because they need to be able to keep an eye out for predators. Interestingly, while they found a relationship between the morphology of the fish and habitat use, they did not find an association between specific feeding tactics and habitat types.

Figure 1. Figure from Lukoschek and McCormick (2001) showing that small fish (black bar) were found in shallow habitat while large fish (white bar) were found in deep habitat.

Conversely, Torres and Read (2009) did find associations between theforaging tactics of bottlenose dolphins in Florida Bay, FL and habitat type. Dolphins in this bay employ three foraging tactics: herd and chase, mud ring feeding, and deep diving. Observations of the foraging tactics were linked to habitat characteristics and individual dolphins. The study found that these tactics are spatially structured by depth (Figure 2), with deep diving occurring in deep water whereas mud ring feeding occurrs in shallower water. They also found evidence of individual specialization! Individuals that were observed deep diving were not observed mud ring feeding and vice-versa. Furthermore, they found that individuals were found in the habitat type associated with their preferred tactic regardless of whether they were foraging or not. This result indicates that individual dolphins in this bay have a foraging tactic they prefer and tend to stay in the corresponding habitat type. These findings are really intriguing and raise interesting questions regarding how these tactics and specializations are developed or learned. These are questions that I am also interested in asking as part of my thesis.

Figure 2. Figure from Torres and Read (2009) showing that deep diving is associated with deeper habitat while mud ring feeding is associated with shallow habitat.

Both of these studies are cool examples that, combined, exemplify questions I am interested in examining using our study population of Pacific Coast Feeding Group (PCFG) gray whales. Like both studies, I am interested in assessing how specific foraging tactics are associated with habitat types. Our hypothesis is that different prey types live in different habitat types, so each tactic corresponds to the best way to feed on that prey type in that habitat. While predation risk doesn’t have as much of an effect on foraging gray whales as it does on small benthic fish, I do wonder how disturbance from boats could similarly affect tactic preference and spatial distribution. I am also curious to see if depth has an effect on tactic choice by using the morphology data from our drone-based photogrammetry. Given that these whales forage in water that is sometimes as deep as they are long, it stands to reason that maneuverability would affect tactic use. As described in a previous blog, I’m also looking for evidence of individual specialization. It will be fascinating to see how foraging preference relates to space use, habitat preference, and morphology.

These studies demonstrate the complexity involved in studying a population’s relationship to its habitat. Such research involves considering the morphology and physiology of the animals, their social, individual, foraging, and predator-prey behaviors, and the relationship between their prey and the habitat. It’s a bit daunting but mostly really exciting because better understanding each puzzle piece improves our ability to estimate how these animals will react to changing environmental conditions.

While I don’t have any answers to these questions yet, I will be working with a National Science Foundation Research Experience for Undergraduates intern this summer to develop a habitat map of our study area that will be used in this analysis and potentially answer some preliminary questions about PCFG gray whale habitat use patterns. So, stay tuned to hear more about our work this summer!


Lukoschek, V., & McCormick, M. (2001). Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): Relationship between foraging behaviour, habitat use, jaw size, and prey selection. Marine Biology, 138(6), 1099–1113.

Torres, L. G., & Read, A. J. (2009). Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins ( Tursiops truncatus ) in Florida Bay, Florida. Marine Mammal Science, 25(4), 797–815.

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

By Solène Derville, Postdoc, OSU Department of Fisheries, Wildlife, and Conservation Science, Geospatial Ecology of Marine Megafauna Lab

The ocean is vast.

What I mean is that the vastness of the ocean is very hard to mentally visualize. When facing a conservation issue such as increased whale entanglement along the US West Coast (see OPAL project ), a tempting solution may  be to suggest « let’s go see where the whales are and report their location to the fishermen?! ». But, it only takes a little calculation to realize how impractical this idea is.

Let’s roll out the numbers. The US West Coast exclusive economic zone (EEZ) stretches from the coast out to 200 nautical miles offshore, as prescribed by the 1982 United Nations Convention on the Law of the Sea. It covers an area of 825,549 km² (Figure 1). Now, imagine that you wish to survey this area for marine mammals. Using a vessel such as the R/V Bell M. Shimada that is used for the Northern California Current Ecosystem surveys cruises (NCC cruises, see Dawn and Rachel’s last blog), we may detect whales at a distance of roughly 6 km (based on my preliminary results). This distance of detection depends on the height of the observer, hence the height of the flying bridge where she/he is standing (the observer’s height may also be accounted for, but unless she/he is a professional basket-ball player, I think it can be neglected here). The Shimada is quite a large ship and it’s flying bridge is 13 meters above the water. Two observers may survey the water on each side of the trackline.

Considering that the vessel is moving at 8 knots (~15 km/h), we may expect to be effectively surveying 180 km² per hour (6x2x15). That’s not too bad, right?

Again, perspective is the key. If we divide the West Coast EEZ surface by 180 km² we can estimate that it would take 2,752 hours to survey this entire region. With an average of 12 hours of daylight, this takes us to…

382 DAYS OF SURVEY, searching for marine mammals over the US West Coast. Considering that observations cannot be undertaken on days with bad weather (fog, heavy rain, strong winds…), it might take more than a year and a half to complete the survey! And what would the marine mammals have done in the meantime? Move…

This little math exercise proves that exhaustively searching for the needle in the haystack from a vessel is not the way to go if we are to describe whale distribution and help mitigate the risk of entanglement. And using another platform of observation is not necessarily the solution. The OPAL project has relied on a great collaboration with the United States Coast Guard to survey Oregon waters. The USCG helicopters travel fast compared to a vessel, about 90 knots (167 km/h). As a result, more ground is covered but the speed at which it is traveling prevents the observer from detecting whales that are very far away. Based on the last analysis I ran for the OPAL project, whales are usually detected up to 3 km from the helicopter (only 5 % of sightings exceed that distance). In addition, the helicopter generally only has capacity for one observer at a time.

If we replicate the survey time calculation from above for the USCG helicopter, we realize that even with a fast-moving aerial survey platform it would still take 137 days to cover the West Coast EEZ.

Figure 1. What is the best survey method to document marine mammal occurrence in the US West Coast Exclusive Economic zone (EEZ)?

First, we can model and extrapolate. This approach is the path we are taking with the OPAL project: we survey Oregon waters in 4 different areas along the coast each month, then model observed whale densities as a function of topographic and oceanographic variables, and then predict whale probability of presence over the entire region. These predictions are based on the assumption that our survey design effectively sampled the variety of environmental conditions experienced by whales over the study region, which it certainly did considering that all sites are surveyed year-round.

An alternative approach that has been recently discussed in the GEMM Llab, is the use of satellite images to detect whales along the coast. A communication entitled « The Potential of Satellite Imagery for Surveying Whales » was published last month in the Sensors Journal (Höschle et al., 2021) and presents the opportunities offered by this relatively new technology. The WorldView-3 satellite, owned by the company Digitalglobe and launched in 2016, has made it possible to commercialize imagery with a resolution never reached before, of the order of 30 cm per pixel. These very high resolution (VHR) satellite images make it possible to identify several species of large whales (Cubaynes et al. al., 2019) and to estimate their density (Bamford et al., 2020). Furthermore, machine learning algorithms, such as Neural Networks, have proved quite efficient at automatically detecting whales in satellite images (Guirado et al., 2019, Figure 2). While several new ultra-high resolution imaging satellites are expected to be launched in 2021 (by Maxar Technologies and Airbus), this “remote” approach looks like a promising avenue to detect whales over vast regions while drinking a cup of coffee at the office.

Figure 2. Illustration of a whale detection algorithm working on a gridded satellite image (DigitalGlobe). Source: Guirado et al., 2019.

But like any other data collection method, satellites have their drawbacks. We recently discovered that these VHR satellites are routinely switched off while passing above the ocean. Specific inquiries would need to be made to acquire data over our study areas, which would be at great expense. One of the cheapest provider I found is the Soar platform, that provides images at 50 cm resolution in partnership with the Chinese Aerospace Science and Technology Corporation. They advertise daily images anywhere on earth at $10 USD per km². This might sound cheap at first glance, but circling back to our US West Coast EEZ area calculations, we estimate that surveying this region entirely with satellite imagery would cost more than $8 million USD.

Yet, we have to look forward. The use of satellite imagery is likely to broaden and increase in the coming years, with a possible decrease in cost. Quoting Höschle et al. (2021) ‘To protect our world’s oceans, we need a global effort and we need to create opportunities for that to happen’.

Will satellites soon save whales?


Bamford, C. C. G. et al. A comparison of baleen whale density estimates derived from overlapping satellite imagery and a shipborne survey. Sci. Rep. 10, 1–12 (2020).

Cubaynes, H. C., Fretwell, P. T., Bamford, C., Gerrish, L. & Jackson, J. A. Whales from space: Four mysticete species described using new VHR satellite imagery. Mar. Mammal Sci. 35, 466–491 (2019).

Guirado, E., Tabik, S., Rivas, M. L., Alcaraz-Segura, D. & Herrera, F. Whale counting in satellite and aerial images with deep learning. Sci. Rep. 9, 1–12 (2019).

Höschle, C., Cubaynes, H. C., Clarke, P. J., Humphries, G. & Borowicz, A. The potential of satellite imagery for surveying whales. Sensors 21, 1–6 (2021).

Defining Behaviors

Clara Bird, PhD Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

When I started working on my thesis, I anticipated many challenges related to studying the behavioral ecology of gray whales. From processing five-plus years of drone footage to data analysis, there has been no shortage of anticipated and unexpected issues. I recently hit an unexpected challenge when I started video processing that piqued my interest. As I’ve discussed in a previous blog, ethograms are lists of defined behaviors that help us properly and consistently collect data in a standardized approach. Ethograms form a crucial foundation of any behavior study as the behaviors defined ultimately affect what questions can be asked and what patterns are detected. Since I am working off of the thorough ethogram of Oregon gray whales from Torres et al. (2018), I had not given much thought to the process of adding behaviors to the ethogram. But, while processing the first chunk of drone videos, I noticed some behaviors that were not in the original ethogram and struggled to decide whether or not to add them. I learned that ethogram development can lead down several rabbit holes. The instinct to try and identify every movement is strong but dangerous. Every minute movement does not necessarily need to be included and it’s important to remember the ultimate goal of the analysis to avoid getting bogged down.

Fundamental behavior questions cannot be answered without ethograms. For example, Baker et al. (2017) developed an ethogram for bottlenose dolphins in Ireland in order to conduct an initial quantitative behavior analysis. They did so by reviewing published ethograms for bottlenose dolphins, consulting with multiple experts, and revising the ethogram throughout the study. They then used their data to test inter-observer variability, calculate activity budgets, and analyze how the activity budgets varied across space and time.

Howe et al. (2015) also developed an ethogram in order to conduct quantitative behavior analyses. Their goals were to use the ethogram and subsequent analyses to better understand the behavior of beluga whales in Cook Inlet, AK, USA and to inform conservation. They started by writing down all behaviors they observed in the field, then they consolidated their notes into a formal ethogram that they used and refined during subsequent field seasons. They used their data to analyze how the frequencies of different behaviors varied throughout the study area at different times. This study served as an initial analysis investigating the effect of anthropogenic disturbance and was refined in future studies.

My research is similarly geared towards understanding behavior patterns to ultimately inform conservation. The primary questions of my thesis involve individual specialization, patterns of behavior across space, the relationship between behavior and body condition, and social behavior (check out this blog to learn more). While deciding what behaviors to add to my ethogram I’ve had to remind myself of these main questions and the bigger picture. The drone footage lets us see so much detail that it’s tempting to try to define every movement we can observe. One rabbit hole I’ve had to avoid a few times is locomotion. From the footage, it is possible to document fluke beats and pectoral fin strokes. While it could be interesting to investigate how different whales move in different ways, it could easily become a complicated mess of classifying different movements and take me deep into the world of whale locomotion. Talking through what that work would look like reminded me that we cannot answer every question and trying to assess all exciting side projects can cause us to lose focus on the main questions.

While I avoided going down the locomotion rabbit hole, there were some new behaviors that I did add to my ethogram. I’ll illustrate the process with the examples of two new behaviors I recently added: fluke swish and pass under (Clips 1 and 2). Clip 1 shows a whale rapidly moving its fluke to the side. I chose to add fluke swish because it’s such a distinct movement and I’m curious to see if there’s a pattern across space, time, individual, or nearby human activity that might explain its function. Clip 2 shows a calf passing under its mom.  It’s not nursing because the calf doesn’t spend time under its mom, it just crosses underneath her. The calf pass under behavior could be a type of mom-calf tactile interaction. Analyzing how the frequency of this behavior changes over time could show how a calf’s dependency on its mom changes over as it ages.

In defining these behaviors, I had to consider how many different variations of this behavior would be included in the definition. This process involves considering at what point a variation of that behavior could serve a different function, even without knowing the function of the original behavior. For fluke swish this process involved deciding to only count a behavior as a fluke swish if it was a big, fast movement. A small and slow movement of the fluke a little to the side could serve a different function, such as turning, or be a random movement.

Clip 1: Fluke swish behavior (Video filmed under NOAA/NMFS research permit #16111​​ by certified drone pilot Todd Chandler).
Clip 2: Pass under behavior (Video filmed under NOAA/NMFS research permit #16111​​ by certified drone pilot Todd Chandler).

The next step involved deciding if the behavior would be a ‘state’ or ‘point’ event. A state event is a behavior with a start and stop moment; a point event is instantaneous and assigned to just a point in time. I would categorize a behavior as a state event if I was interested in questions about its duration. For example, I could ask “what percentage of the total observation time was spent in a certain behavior state?” A point event would be a behavior where duration is not applicable, but I could ask a question like “Did whale 1 perform more point event A than whale 2?”. Both fluke swish and pass under are point events because they only happen for an instant. In a pass under the calf is passing under its mom for just a brief point in time, making it a point event. The final step was to name the behavior. As I discussed in this blog, the name of the behavior does not matter as much as the definition but it is important that the name is clear and descriptive. We chose the name fluke swish because the fluke rapidly moves from side to side and pass under because the calf crosses under its mom.

Frankly, in the beginning, I was a bit overwhelmed by the realization that the content of my ethogram would ultimately control the questions I could answer. I could not help but worry that after processing all the videos, I would end up regretting not defining more behaviors. However, after reading some of the literature, chatting with Leigh, and reviewing the initial chunk of videos several times, I am more confidence in my judgment and my ethogram. I have accepted the fact that I can’t anticipate everything, and I am confident that the behaviors I need to answer my research questions are included. The process of reviewing and updating my ethogram has been a rewarding challenge that resulted in a valuable lesson that I will take with me for the rest of my career.


Baker, I., O’Brien, J., McHugh, K., & Berrow, S. (2017). An ethogram for bottlenose dolphins (Tursiops truncatus) in the Shannon Estuary, Ireland. Aquatic Mammals, 43(6), 594–613.

Howe, M., Castellote, M., Garner, C., McKee, P., Small, R. J., & Hobbs, R. (2015). Beluga, Delphinapterus leucas, ethogram: A tool for cook inlet beluga conservation? Marine Fisheries Review, 77(1), 32–40.

Torres, L. G., Nieukirk, S. L., Lemos, L., & Chandler, T. E. (2018). Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science, 5(SEP).

Lessons learned from (not) going to sea

By Rachel Kaplan1 and Dawn Barlow2

1PhD student, Oregon State University College of Earth, Ocean, and Atmospheric Sciences and Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

2PhD Candidate, Oregon State University Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

“Hurry up and wait.” A familiar phrase to anyone who has conducted field research. A flurry of preparations, followed by a waiting game—waiting for the weather, waiting for the right conditions, waiting for unforeseen hiccups to be resolved. We do our best to minimize unknowns and unexpected challenges, but there is always uncertainty associated with any endeavor to collect data at sea. We cannot control the whims of the ocean; only respond as best we can.

On 15 February 2021, we were scheduled to board the NOAA Ship Bell M. Shimada as marine mammal observers for the Northern California Current (NCC) ecosystem survey, a recurring research cruise that takes place several times each year. The GEMM Lab has participated in this multidisciplinary data collection effort since 2018, and we are amassing a rich dataset of marine mammal distribution in the region that is incorporated into the OPAL project. February is the middle of wintertime in the North Pacific, making survey conditions challenging. For an illustration of this, look no further than at the distribution of sightings made during the February 2018 cruise (Fig. 1), when rough sea conditions meant only a few whales were spotted.

Figure 1. (A) Map of marine mammal survey effort (gray tracklines) and baleen whale sightings recorded onboard the NOAA ship R/V Shimada during each of the NCC research cruises to-date and (B) number of individuals sighted per cruise since 2018. Note the amount of survey effort conducted in February 2018 (top left panel) compared to the very low number of whales sighted. Data summary and figures courtesy of Solene Derville.

Now, this is February 2021 and the world is still in the midst of navigating the global coronavirus pandemic that has affected every aspect of our lives. The September 2020 NCC cruise was the first NOAA fisheries cruise to set sail since the pandemic began, and all scientists and crew followed a strict shelter-in-place protocol among other COVID risk mitigation measures. Similarly, we sheltered in place in preparation for the February 2021 cruise. But here’s where the weather comes in yet again. Not only did we have to worry about winter weather at sea, but the inclement conditions across the country meant our COVID tests were delayed in transit—and we could not board the ship until everyone tested negative. By the time our results were in, the marine forecast was foreboding, and the Captain determined that the weather window for our planned return to port had closed.

So, we are still on shore. The ship never left the dock, and NCC February 2021 will go on the record as “NAs” rather than sightings of marine mammal presence or absence. So it goes. We can dedicate all our energy to studying the ocean and these spectacularly dynamic systems, but we cannot control them. It is an important and humbling reminder. But as we have continued to learn over the past year, there are always silver linings to be found.

Even though we never made it to the ship, it turns out there’s a lot you can get done onshore. Dawn has sailed on several NCC cruises before, and one of the goals this time was to train Rachel for her first stint at marine mammal survey work. This began at Dawn’s house in Newport, where we sheltered in place together for the week prior to our departure date.

We walked through the iPad program we use to enter data, looked through field guides, and talked over how to respond in different scenarios we might encounter while surveying for marine mammals at sea. We also joined Solene, a postdoc working on the OPAL project, for a Zoom meeting to edit the distance sampling protocol document. It was great training to discuss the finer points of data collection together, with respect to how that data will ultimately be worked into our species distribution models.

The February NCC cruise is famously rough, and a tough time to sight whales (Fig. 1). This low sighting rate arises from a combination of factors: baleen whales typically spend the winter months on their breeding grounds in lower latitudes so their density in Oregon waters is lower, and the notorious winter sea state makes sighting conditions difficult. Solene signed off our Zoom call with, “Go collect that high-quality absence data, girls!” It was a good reminder that not seeing whales is just as important scientifically as seeing them—though sometimes, of course, it’s not possible to even get out where you can’t see them. Furthermore, all absence data is not created equal. The quality of the absence data we can collect deteriorates along with the weather conditions. When we ultimately use these survey data to fuel species distribution models, it’s important to account for our confidence in the periods with no whale sightings.

In addition to the training we were able to conduct on land, the biggest silver lining came just from sheltering in place together. We had only met over Zoom previously, and spending this time together gave us the opportunity to get to know each other in real life and become friends. The week involved a lot of fabulous cooking, rainy walks, and an ungodly number of peanut butter cups. Even though the cruise couldn’t happen, it was such a rich week. The NCC cruises take place several times each year, and the next one is scheduled for May 2021. We’ll keep our fingers crossed for fair winds and negative COVID tests in May!

Figure 2. Dawn’s dog Quin was a great shelter in place buddy. She was not sad that the cruise was canceled.

The ups and downs of the ocean

By Solène Derville, Postdoc, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

As a GEMM lab post-doc working on the OPAL project, my main goal for 2021 will be to produce accurate predictive models of baleen whale distribution off the Oregon coast to reduce entanglement risk. For the past months, I have been compiling, cleaning, and processing about two years of data collected by Leigh Torres and Craig Hayslip during monthly repeat surveys conducted onboard United States Coast Guard (USCG) helicopters. These standardized surveys record where and when whales are observed off the Oregon coast. These presence and absence data may now be modeled in relation to habitat, while accounting for effort and detection (as several parameters, such as weather and sea state, can affect the capacity of observers to detect whales at the surface). Considering that several baleen whale species (namely, humpback, fin, blue and gray whales) are known to feed in the area, prey availability is expected to be a major driver of their distribution.

As prey distribution data are frequently the lacking component in the habitat model equation, whale ecologists often resort to using environmental proxies. Variables such as topography (e.g., the depth or slope of the seafloor), water physical and chemical characteristics (e.g., temperature, salinity, oxygen concentration) or ocean circulation (e.g., currents, turbulence) have proved to be good predictors for fish or krill distribution, and in turn potential predictors for whale suitable habitats. In my search for such environmental variables to be tested in our future OPAL models, I have been focusing my research on a fascinating ocean feature: sea height.

Sea height varies both temporally and spatially under the influence of multiple factors, from internal mass of the solid Earth to the orbital revolution of the moon. After reading this blog you will realize that the flatness of the horizon at sea is a deceiving perspective (Figure 1) …

Figure 1: Flat? Really? (source: Pixabay)

Gravity and the geoid

We all know of Newton’s s discovery of gravity: the attraction force exerted by any object with a given mass on its surroundings. Yet, it is puzzling to think that the rate of acceleration of the apple falling on Newton’s head would have been different if Newton had been anywhere else on Earth.

Why is that and what does it have to do with sea height? On Earth, the standard gravity g is set at 9.80665 m/s2. This constant is called a “standard” because in fact, gravity varies at the surface of our planet, even if estimated at a fixed altitude. Indeed, as gravity is caused by mass, any change in relief or rock composition results in a change in gravity. For instance, magmatic activity in the upper mantle of the Earth and the crust causes a change in rock density and results in a change in gravity measured at the surface.

Gravity therefore is the first reason why the ocean surface is not flat. Gravity shapes an irregular surface called the “geoid”. This hypothetical ocean surface has equal gravitational potential anywhere on Earth and differs from the ellipsoid of reference by as much as 100 m! So to the question whether Earth is round or flat, I would say it is potato shaped (Figure 2)!

Figure 2: Exaggerated view of the gravitational potential of Earth. View a video animation here. (credit: European Space Agency)

The geoid is an essential reference for understanding ocean currents and monitoring changes in sea-level. Hypothetically, if ocean water had equal density everywhere and at any depth, the sea surface should match with the geoid… but that’s not the case. Let’s see why.

Ocean dynamic topography

Not unlike the hills and valleys covering landscapes, the ocean surface also has its highs and lows. Except that in the ocean, the surface topography is ever changing. Sea surface height (SSH) measures the average height difference between the observed sea level and the ellipsoid of reference (Figure 3). SSH is mostly affected by ocean circulation and may vary by as much as ±1 m. Indeed, just like the rocks inside the Earth, the water in the ocean varies in density. The vertical and horizontal physical structuring of the ocean was extensively discussed by Dawn last November while she was preparing for her PhD Qualifying Exams. Temperature clearly is at the core of the processes. As thermal expansion increases the space between warming water particles, the volume of a given amount of liquid water increases with increasing temperature. Warmer waters therefore take up “more space” than cooler waters, resulting in an elevated SSH.

Figure 3: Overview of the different fields used in altimetry (credit: CLS,

SSH may therefore be used as an indicator of oceanographic phenomena such as upwellings, where warm surface waters are replaced by deep, cooler, and nutrient-rich waters moving upwards. The California Current that moves southwards along the North American coast is known as one of the world’s major currents affiliated with strong upwelling zones, which often triggers increased biological productivity. Several studies conducted in the California Current system have found a link between the variations in SSH and whale abundance or foraging activity (Abrahms et al. 2019; Pardo et al. 2015; Becker et al. 2016; Hazen et al. 2016).⁠

SSH is measured by altimeter satellites and is made freely available by the European Space Agency and the US National Aeronautics and Space Administration. Lucky me! Numerous variables are derived from SSH, as shown in Figure 3. Among other things, I was able to download the daily maps of Sea Surface Height Anomaly (SSHa, also referred to as Sea Level Anomaly: SLA) over the Oregon coast from February 2019 to December 2020. SSHa is the difference between observed SSH at a specific time and place from the mean SSH field of reference calculated over a long period of time. Negative values of SSHa potentially suggest upwellings of cooler waters that could be associated with higher prey availability. Figure 4 shows an example of environmental data mining as I try to match SSHa with whale observations made during OPAL surveys. Figure 4B suggests increased whale occurrence where/when SSHa is lower.

Figure 4: Preliminary exploration of the relationship between sea surface height anomaly (SSHa) and baleen whales (blue, fin, humpback, unidentified) observed during OPAL surveys off Oregon, USA, between February 2019 and December 2020. A) Example covering 3 months of survey during summer 2019. Sightings were grouped over 5-km segments of surveyed trackline and segments with at least one sighting were mapped with colored circles. Dotted grey lines are the repeated survey tracklines for each of the labeled study areas (NB = North Bend). Sightings are symbolized by area (color)
and group size (circle size). Monthly averages of SSHa are represented with a colored gradient. B) Monthly averages of SSHa measured over 5-km segments where whales were detected (presence) or not (absence).

Although encouraging, these preliminary insights are just the tip of the modeling iceberg. Many more testing and modeling steps will be required to determine confounding factors and relevant spatio-temporal scales at which these oceanographic variables may be influencing whale distribution off the Oregon coast. I am only at the start of a long road…


Abrahms, Briana, Heather Welch, Stephanie Brodie, Michael G. Jacox, Elizabeth A. Becker, Steven J. Bograd, Ladd M. Irvine, Daniel M. Palacios, Bruce R. Mate, and Elliott L. Hazen. 2019. “Dynamic Ensemble Models to Predict Distributions and Anthropogenic Risk Exposure for Highly Mobile Species.” Diversity and Distributions, no. December 2018: 1–12.

Becker, Elizabeth, Karin Forney, Paul Fiedler, Jay Barlow, Susan Chivers, Christopher Edwards, Andrew Moore, and Jessica Redfern. 2016. “Moving Towards Dynamic Ocean Management: How Well Do Modeled Ocean Products Predict Species Distributions?” Remote Sensing 8 (2): 149.

Hazen, Elliott L, Daniel M Palacios, Karin A Forney, Evan A Howell, Elizabeth Becker, Aimee L Hoover, Ladd Irvine, et al. 2016. “WhaleWatch : A Dynamic Management Tool for Predicting Blue Whale Density in the California Current.” Journal of Applied Ecology 54 (5): 1415–28.

Pardo, Mario A., Tim Gerrodette, Emilio Beier, Diane Gendron, Karin A. Forney, Susan J. Chivers, Jay Barlow, and Daniel M. Palacios. 2015. “Inferring Cetacean Population Densities from the Absolute Dynamic Topography of the Ocean in a Hierarchical Bayesian Framework.” PLOS One 10 (3): 1–23.

Are there picky eaters in the PCFG?

Clara Bird, PhD Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

As anyone who has ever been, or raised, a picky eater knows, humans have a wide range of food preferences. The diversity of available cuisines is a testament to the fact that we have individual food preferences. While taste is certainly a primary influence, nutritional benefits and accessibility are other major factors that affect our eating choices. But we are not the only species to have food preferences. In cetacean research, it is common to study the prey types consumed by a population as a whole. Narrowing these prey preferences down to the individual level is rare. While the individual component is challenging to study and to incorporate into population models, it is important to consider what the effects of individual foraging specialization might be.

To understand the role and drivers of individual specialization in population ecology, it is important to first understand the concepts of niche variation and partitioning. An animal’s ecological niche describes its role in the ecosystem it inhabits (Hutchinson, 1957). A niche is multidimensional, with dimensions for different environmental conditions and resources that a species requires. One focus of my research pertains to the dimensions of the niche related to foraging. As discussed in a previous blog, niche partitioning occurs when ecological space is shared between competitors through access to resources varies across different dimensions such as prey type, foraging location, and time of day when foraging takes place. Niche partitioning is usually discussed on the scale of different species coexisting in an ecosystem. Pianka’s theory stating that niche partitioning will increase as prey availability decreases uses competing lizard species as the example (Pianka, 1974). Typically, niche partitioning theory considers inter-specific competition (competition between species), but niche partitioning can take place within a species in response to intra-specific competition (competition between individuals of the same species) through individual niche variation.

A species that consumes a multitude of prey types is considered a generalist while one with a specific prey type is considered a specialist. Gray whales are considered generalists (Nerini, 1984). However, we do not know if each individual gray whale is a generalist or if the generalist population is actually composed of individual specialists with different preferences. One way to test for the presence of individual specialization is to compare the niche width of the population to the niche width of each individual (Figure 1, Bolnick et al., 2003).  For example, if a population eats five different types of prey and each individual consumed those prey types, those individuals would be generalists. However, if each individual only consumed one of the prey types, then those individuals would be specialists within a generalist population.

Figure 1. Figure from Bolnick et al. 2003. The thick curve represents the total niche of the population and the thin curves represent individual niches. Note that in both panels the population has the same total niche. In panel A, the individual curves overlap and are all pretty wide. These curves represent individual generalists that make up a generalist population. In panel B, the thin curves are narrower and do not overlap as much as those in panel A. These curves represent individual specialists that make up a generalist population.

If individual specialization is present in a population the natural follow-up question is why? To answer this, we look for common characteristics between the individuals that are similarly specialized. What do all the individuals that feed on the same prey type have in common? Common characterizations that may be found include age, sex, or distinct morphology (such as different beak or body shapes) (Bolnick et al., 2003).

Woo et al. (2008) studied individual specialization in Brünnich’s guillemot, a generalist sea bird species, using diet and tagging data. They found individual specialization in both diet (prey type) and behavior (dive depth, shape, and flight time). Specialization occurred across multiple timescales but was higher over short-time scales. The authors found that it was more common for an individual to specialize in a prey-type/foraging tactic for a few days than for that specialization to continue across years, although a few individuals were specialists for the full 15-year period of the study. Based on reproductive success of the studies birds, the authors concluded that the generalist and specialist strategies were largely equivalent in terms of fitness and survival. The authors searched for common characteristics in the individuals with similar specialization and they found that the differences between sexes or age classes were so small that neither grouping explained the observed individual specialization. This is an interesting result because it suggests that there is some missing attribute, that of the authors did not examine, that might explain why individual specialists were present in the population.

Hoelzel et al. (1989) studied minke whale foraging specialization by observing the foraging behaviors of 23 minke whales over five years from a small boat. They identified two foraging tactics: lunge feeding and bird-associated feeding. Lunge feeding involved lunging up through the water with an open mouth to engulf a group of fish, while bird-associated feeding took advantage of a group of fish being preyed on by sea birds to attack the fish from below while they were already being attacked from above. They found that nine individuals used lunge feeding, and of those nine, six whales used this tactic exclusively. Five of those six whales were observed in at least two years. Seventeen whales were observed using bird-associated feeding, 14 exclusively. Of those 14, eight were observed in at least two years. Interestingly, like Woo et al. (2008), this study did not find any associations between foraging tactic use and sex, age, or size of whale. Through a comparison of dive durations and feeding rates, they hypothesized that lunge feeding was more energetically costly but resulted in more food, while bird-associated feeding was energetically cheaper but had a lower capture rate. This result means that these two strategies might have the similar energetic payoffs.

Both of these studies are examples of questions that I am excited to ask using our data on the PCFG gray whales feeding off the Oregon coast (especially after doing the research for this blog). We have excellent individual-specific data to address questions of specialization because the field teams for  this project always carefully link observed behaviors with individual whale ID.  Using these data, I am curious to find out if the whales in our study group are individual specialists or generalists (or some combination of the two). I am also interested in relating specific tactics to their energetic costs and benefits in order to assess the payoffs of each foraging tactic. I then hope to combine the results of both analyses to assess the payoffs of each individual whale’s strategy.

Figure 2. Example images of two foraging tactics, side swimming (left) and headstanding (right). Images captured under NOAA/NMFS permit #21678.

Studying individual specialization is important for conservation. Consider the earlier example of a generalist population that consumes five prey items but is composed of individual specialists. If the presence of individual specialization is not accounted for in management plans, then regulations may protect certain prey types or foraging tactics/areas of the whales and not others. Such a management plan could be a dangerous outcome for the whale population because only parts of the population would be protected, while other specialists are at risk, thus potentially losing genetic diversity, cultural behaviors, and ecological resilience in the population as a whole. A plan designed to maximize protection for all the specialists would be better for the population because populations with increased ecological resilience are more likely to persist through periods of rapid environmental change. Furthermore, understanding individual specialization could help us better predict how a population might be affected by environmental change. Environmental change does not affect all prey species in the same way. An individual specialization study could help identify which whales might be most affected by predicted environmental changes. Therefore, in addition to being a fascinating and exciting research question, it is important to test for individual specialization in order to improve management and our overall understanding of the PCFG gray whale population.


Bolnick, D. I., Svanbäck, R., Fordyce, J. A., Yang, L. H., Davis, J. M., Hulsey, C. D., & Forister, M. L. (2003). The ecology of individuals: Incidence and implications of individual specialization. American Naturalist, 161(1), 1–28.

Hoelzel, A. R., Dorsey, E. M., & Stern, S. J. (1989). The foraging specializations of individual minke whales. Animal Behaviour, 38(5), 786–794.

Hutchinson, G. E. (1957). Concluding Remarks. Cold Spring Harbor Symposia on Quantitative Biology, 22(0), 415–427.

Nerini, M. (1984). A Review of Gray Whale Feeding Ecology. In The Gray Whale: Eschrichtius Robustus (pp. 423–450). Elsevier Inc.

Pianka, E. R. (1974). Niche Overlap and Diffuse Competition. 71(5), 2141–2145.

Woo, K. J., Elliott, K. H., Davidson, M., Gaston, A. J., & Davoren, G. K. (2008). Individual specialization in diet by a generalist marine predator reflects specialization in foraging behaviour. Journal of Animal Ecology, 77(6), 1082–1091.