Tag: PCFG

A smaller sized gray whale: recent publication finds PCFG whales are smaller than ENP whales

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

A recent blog post by GEMM Lab’s PhD Candidate Clara Bird gave a recap of our 8th consecutive GRANITEfield season this year. In her blog, Clara highlighted that we saw 71 individual gray whales this season, 61 of which we have seen in previous years and identified as belonging to the Pacific Coast Feeding Group (PCFG). With an estimated population size of around 212 individuals, this means that we saw almost 1/3 of the PCFG population this season alone. Since the GEMM Lab first started collecting data on PCFG gray whales in 2016, we have collected drone imagery on over 120 individuals, which is over half the PCFG population. This dataset provides incredible opportunity to get to know these individuals and observe them from year to year as they grow and mature through different life history stages, such as producing a calf. A question our research team has been interested in is what makes a PCFG whale different from an Eastern North Pacific (ENP) gray whale, which has a population size around 16,000 individuals and feed predominantly in the Arctic during the summer months? For this blog, I will highlight findings from our recent publication in Biology Letters (Bierlich et al., 2023) comparing the morphology (body length, skull, and fluke size) between PCFG and ENP populations. 

Body size and shape reflect how an animal functions in their environment and can provide details on an individual’s current health, reproductive status, and energetic requirements. Understanding how animals grow is a key component for monitoring the health of populations and their vulnerability to climate change and other stressors in their environment.  As such, collecting accurate morphological measurements of individuals is essential to model growth and infer their health. Collecting such morphological measurements of whales is challenging, as you cannot ask a whale to hold still while you prepare the tape measure, but as discussed in a previous blog, drones provide a non-invasive method to collect body size measurements of whales. Photogrammetry is a non-invasive technique used to obtain morphological measurements of animals from photographs. The GEMM Lab uses drone-based photogrammetry to obtain morphological measurements of PCFG gray whales, such as their body length, skull length (as snout-to-blowhole), and fluke span (see Figure 1). 

Figure 1. Morphological measurements obtained via photogrammetry of a Pacific Coast Feeding Group (PCFG) gray whale. These measurements were used to compare to individuals from the Eastern North Pacific (ENP) population. 

As mentioned in this previous blog, we use photo-identification to identify unique individual gray whales based on markings on their body. This method is helpful for linking all the data we are collecting (morphology, hormones, behavior, new scarring and skin conditions, etc.) to each individual whale. An individual’s sightings history can also be used to estimate their age, either as a ‘minimum age’ based on the date of first sighting or a ‘known age’ if the individual was seen as a calf. By combining the length measurements from drone-based photogrammetry and age estimates from photo-identification history, we can construct length-at-age growth models to examine how PCFG gray whales grow. While no study has previously examined length-at-age growth models specifically for PCFG gray whales, another study constructed growth curves for ENP gray whales using body length and age estimates obtained from whaling, strandings, and aerial photogrammetry (Agbayani et al., 2020). For our study, we utilized these datasets and compared length-at-age growth, snout-to-blowhole length, and fluke span between PCFG and ENP whales. We used Bayesian statistics to account and incorporate the various levels of uncertainty associated with data collected (i.e., measurements from whaling vs. drone, ‘minimum age’ vs. ‘known age’). 

We found that while both populations grow at similar rates, PCFG gray whales reach smaller adult lengths than ENP. This difference was more extreme for females, where PCFG females were ~1 m (~3 ft) shorter than ENP females and PCFG males were ~0.5 m (1.5 ft) shorter than ENP males (Figure 2, Figure 3). We also found that ENP males and females have slightly larger skulls and flukes than PCFG male and females, respectively. Our results suggest PCFG whales are shaped differently than ENP whales (Figure 3)! These results are also interesting in light of our previous published study that found PCFG whales are skinnier than ENP whales (see this previous blog post). 

Figure 2. Growth curves (von Bertalanffy–Putter) for length-at-age comparing male and female ENP and PCFG gray whales (shading represents 95% highest posterior density intervals). Points represent mean length and median age. Vertical bars represent photogrammetric uncertainty. Dashed horizontal lines represent uncertainty in age estimates.

Figure 3. Schematic highlighting the differences in body size between Pacific Coast Feeding Group (PCFG) and Eastern North Pacific (ENP) gray whales. 

Our results raise some interesting questions regarding why PCFG are smaller: Is this difference in size and shape normal for this population and are they healthy? Or is this difference a sign that they are stressed, unhealthy and/or not getting enough to eat? Larger individuals are typically found at higher latitudes (this pattern is called Bergmann’s Rule), which could explain why ENP whales are larger since they feed in the Arctic. Yet many species, including fish, birds, reptiles, and mammals, have experienced reductions in body size due to changes in habitat and anthropogenic stressors (Gardner et al., 2011). The PCFG range is within closer proximity to major population centers compared to the ENP foraging grounds in the Arctic, which could plausibly cause increased stress levels, leading to decreased growth. 

The smaller morphology of PCFG may also be related to the different foraging tactics they employ on different prey and habitat types than ENP whales. Animal morphology is linked to behavior and habitat (see this blogpost). ENP whales feeding in the Arctic generally forage on benthic amphipods, while PCFG whales switch between benthic, epibenthic and planktonic prey, but mostly target epibenthic mysids. Within the PCFG range, gray whales often forage in rocky kelp beds close to shore in shallow water depths (approx. 10 m) that are on average four times shallower than whales feeding in the Arctic. The prey in the PCFG range is also found to be of equal or higher caloric value than prey in the Arctic range (see this blog), which is interesting since PCFG were found to be skinnier.

It is also unclear when the PCFG formed? ENP and PCFG whales are genetically similar, but photo-identification history reveals that calves born into the PCFG usually return to forage in this PCFG range, suggesting matrilineal site fidelity that contributes to the population structure. PCFG whales were first documented off our Oregon Coast in the 1970s (Figure 4). Though, from examining old whaling records, there may have been PCFG gray whales foraging off the coasts of Northern California to British Columbia since the 1920s.

Figure 4. First reports of summer-resident gray whales along the Oregon coast, likely part of the Pacific Coast Feeding Group. Capital Journal, August 9, 1976, pg. 2.

Altogether, our finding led us to two hypotheses: 1) the PCFG range provides an ecological opportunity for smaller whales to feed on a different prey type in a shallow environment, or 2) the PCFG range is an ecological trap, where individuals gain less energy due to energetically costly feeding behaviors in complex habitat while potentially targeting lower density prey, causing them to be skinnier and have decreased growth. Key questions remain for our research team regarding potential consequences of the smaller sized PCFG whales, such as does the smaller body size equate to reduced resilience to environmental and anthropogenic stressors? Does smaller size effect fecundity and population fitness? Stay tuned as we learn more about this unique and fascinating smaller sized gray whale. 

References

Agbayani, S., Fortune, S. M. E., & Trites, A. W. (2020). Growth and development of North Pacific gray whales (Eschrichtius robustus). Journal of Mammalogy, 101(3), 742–754. https://doi.org/10.1093/jmammal/gyaa028

Bierlich, K. C., Kane, A., Hildebrand, L., Bird, C. N., Fernandez Ajo, A., Stewart, J. D., Hewitt, J., Hildebrand, I., Sumich, J., & Torres, L. G. (2023). Downsized: gray whales using an alternative foraging ground have smaller morphology. Biology Letters, 19(8). https://doi.org/10.1098/rsbl.2023.0043

Gardner, J. L., Peters, A., Kearney, M. R., Joseph, L., & Heinsohn, R. (2011). Declining body size: A third universal response to warming? Trends in Ecology and Evolution, 26(6), 285–291. https://doi.org/10.1016/j.tree.2011.03.005

Introducing IndividuWhale!

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

If you are an avid reader of our blog, you probably know quite a bit about gray whales, specifically the Pacific Coast Feeding Group (PCFG) of gray whales. Of the just over 50 GEMM Lab blogs written in 2021, 43% of them were about PCFG gray whales (or at least mentioned gray whales in some way). I guess this statistic is not too surprising when you consider that six of the 10 GEMM Lab members conduct gray whale-related research. You might think that we would have reached our annual limit of online gray whale content with that many blogs featuring these gentle giants, but you would in fact be wrong…

At the end of 2021, we launched a brand new website all about gray whales called IndividuWhale! It features stories of some of the Oregon coast’s most iconic gray whales, as well as information about how we study them, stressors they experience in our waters, and even a game to test your gray whale identification skills. IndividuWhale is a true labor of love that took over a year to create and that we are extremely proud to share with you today. Before I tell you more about the website, I want to take a moment to give a huge shout out to Erik Urdahl who was instrumental in getting this website off the ground and making it as interactive and beautiful as it is – hurrah Erik!

Equal‘s right side with visible boat propeller scars. Source: GEMM Lab.

Like us humans, gray whales have individual personalities and stories. They experience life-altering events, go through periods of stress, must provide for their offspring, and can behave differently to one another. Since Leigh & co. have been conducting in-depth research about PCFG gray whales in Oregon waters since 2016, we have been able to document several fascinating stories and events that these individuals have experienced. Take Equal, for example, a male whale that is at least 7 years old. The GEMM Lab observed Equal on consecutive days in June 2018, where on the first day he looked healthy and normal, but on the second day had fresh boat propeller-like scars on his back. Not only did we document these scars in photographs, but we were also able to collect a fecal sample from Equal the day we observed him with these scars. After analyzing his fecal sample for stress hormones, we discovered that Equal had very high stress levels compared to previous samples collected – unsurprising seeing as he had been hit by a boat! While this event was certainly sad for Equal (although don’t worry – we have seen him many times again in the years after this event looking healthy & normal once again), it was a very fortuitous occurrence for us since we were able to “validate” our stress hormone data relative to the value from Equal when he was clearly stressed out. Find out more about Equal as well as seven other gray whales here!

You might be wondering, how we knew that the whale with the boat propeller scar was Equal and how we recognize him again years after the incident. Gray whales have unique pigmentation patterns on their bodies and flukes that allow us to re-identify individuals between years and distinguish them from one another. Additionally, scars, such as those that Equal now carries on his back, can also be useful in telling whales apart. Therefore, we take photographs of every whale we see to match markings and identify whales. This process is called photo ID. Some individuals can have very distinctive markings, such as Roller Skate who has two big white dots on her right side, while others can look more inconspicuous, like Clouds. Therefore, conducting photo ID requires a lot of attention to detail and perseverance. To learn more about the different features we use to identify individuals, check out the “Studying Whales With Photographs” page. Do you think you have what it takes to tell individuals apart? Then try your luck at our photo ID game after!

Test your photo ID skills in our whale match game!

Unfortunately, these whales do not live in a pristine environment, as is evidenced by Equal’s story. Their main objective during the summer when in Oregon waters is to gain weight (energy stores) by consuming large amounts of prey, which is made more difficult by a number of stressors, including potential fishery entanglements, ocean noise, vessel traffic, and habitat changes. We describe these four stressors on the IndividuWhale website since we are trying to assess the impacts of them on gray whales through our research, however they are certainly not the only stressors that these whales experience. Little is known about the level at which these stressors may have a negative impact on the whales, and how whales react when they experience some of these stressors in concert. The answers to these questions are difficult to tease apart but the GRANITE project is aiming to do so through a framework called Population Consequences of Multiple Stressors (read more about it here). This approach requires a lot of data on a lot of individuals in a population and as you can see from the IndividuWhale website, we are slowly starting to get there! 2022 will certainly bring many more gray whale-themed blogs to this website, and you can share in our journey of learning about the lives of PCFG gray whales by exploring the IndividuWhale website (https://www.individuwhale.com).

Stories from a whale’s life: how whale photo-identification is an essential part of the whale physiology toolkit

By Alejandro Fernandez Ajo, PhD student at the Department of Biology, Northern Arizona University, Visiting scientist in the GEMM Lab working on the gray whale physiology and ecology project  

Two years ago, in August 2018, I came to Newport and visited the Hatfield Marine Science Center for the first time with an NSF/RCN-founded laboratory exchange with the GEMM Lab and met Dr. Leigh Torres. My goals during this exchange where to learn about non-invasive fieldwork techniques for studying free-range whales while interacting, exchanging ideas, and networking with the GEMM Lab members; also, to discuss some projects and thoughts for future collaborations with Dr. Torres. During those two weeks in Newport, I had the opportunity to help with field work on the project “Evaluation of gray whale ecology and physiology in response to variable ambient ocean noise conditions”, which aims to evaluate the hormonal variability and health of the gray whales that forage along the Oregon coast in the context of multiple stressors. I would return during the summers of 2019 and 2020 as a visiting scientist and research assistant to work on this project. This year the experience has been a bit different in terms of interactions with the HMSC community due to COVID-19; however, we were able to successfully start the field season in time and now we are wrapping up our second month of surveys with many new and interesting data gathered, and many new, unforgettable memories to be treasured. Working with these animals is incredibly fascinating because there are so many things we don´t know about them, and the questions can become both overwhelming and exciting.

An essential part of this project, and arguably any research project done with cetaceans, is the identification of individuals. Hence, considerable effort is expended each year attempting to photograph every gray whale possible within our study region and to identify each whale we encounter. The GEMM Lab maintains a catalog of the gray whales that visit the Oregon coast, a sub-population known as the Pacific Coast Feeding Group (PCFG). This catalog currently consists of 173 individuals. which we frequently compare with a larger catalog of gray whales that includes 2060 individuals observed since 1977 (Cascadia Research Collective). These methods allow us to know who is who among the whales we encounter each day at sea.

The different species of cetaceans can be individually identified by markings on their bodies, very much like fingerprints in humans. Some features on these animals are unique and conserved through life. For example, Southern and Northern right whales are identified by the callosity patters in their heads (Picture 1), while humpback whales are mostly identified individually by the shape and the patterns of black and white pigmentation on the underside of their fluke (Picture 2). Gray whales have very mottled skin coloration, so we use a combination of markings and features to identify individuals: pigmentation patterns, scars, shape and pigmentation of their fluke, and sometimes the shape of their knuckles, which are a series of “humps” that gray whales have instead of a dorsal fin on their back. It might sound very difficult to do, and it can be a tedious task, however as you train your eye it becomes easier, and features that at first seemed undistinguishable become recognizable and unique (Picture 3). As a reward, it is such a joy to find a match and recognize old friends when they arrive from their long journeys in the vast ocean each year to the Oregon coast.

The callosity pattern of a southern right whale. Photo by Alejandro Fernández Ajó – Instituto de Conservación de Ballenas.
Three different humpback whales with notably different pigmentation patterns on their flukes. Photographs captured under NOAA/NMFS permit #21678.
The gray whale “Knife” we have observed frequently this summer off the Oregon coast. Can you see how the right pigmentation pattern identified looks like a knife? Photograph captured under NOAA/NMFS permit #21678.

As a result of our photo-identification efforts and the high site-fidelity of the whales we study, the large majority of the gray whales we observe here in Oregon are known individually. For many whales, we also have detailed sightings records that can span years and decades, that document calving history, lactation, appearance of scars indicative of injury or entanglement, minimum age, sex, habitat-use patterns, behaviors, etc. Holding such detailed information of individual whales provides incredible contributions to our understanding of the basic patterns in life history of whales, such as reproduction rates, calving intervals, age of first reproduction, etc. Moreover, when these life history events are linked with physiological sample collection large steps can be made in the development and validation of physiological methods. Many endocrine assays currently in use for whales are based on non-traditional sample types including feces, respiratory vapor, and baleen, which have been validated using the catalogs of well-known individuals to verify that measured hormones reflect patterns expected for various physiological states. For example, we can compare endocrine data of confirmed pregnant females, known mature males, and known-injured whales to learn how whale physiological responses are different during different life history events (e.g., Burgess et al. 2017, 2018, Corkeron et al. 2017, Hunt et al. 2006, 2016, 2018, Lysiak et al. 2018, Rolland et al. 2005).

Here in Oregon we are learning from the lives of the gray whales we study, and here I want to share with you two of their stories, one happy and one not-so-happy.

Let´s start with the not-so-happy story so we can end with some brighter news. On June 24 this year, we encountered a whale near Cape Foulweather, which is a very tricky area to work as there are many rocks and shallow water that make the sea conditions very choppy even with low swell. We started documenting the sighting as usual, taking pictures of the left side, the right side and ideally also the fluke of the whale. As we approached this whale, we started noting that something was wrong with its fluke. With the challenging sea conditions, it was not easy to approach the whale and the whale was not exposing its fluke when diving. When we put our drone up to collect photogrammetry and behavior data we gained a much better perspective. This whale has a bad injury on it fluke (Picture 4.C). On the boat we started making conjectures about the cause of this terrible injury that had basically amputate most of its left fluke lobe. Once back on shore, we sorted out the photos and compared the field images captured during the day with the photo-ID catalog and we made a match. This whale is known in our catalog as “ROLLER SKATE”, is a female, and was first sighted in 2015, so she is at least 5 years old today.

The story unfolded when we reviewed Roller Skate’s sighting history. Interestingly we observed this same whale in the same location last September 2019. Unfortunately, it was a very brief encounter but enough for photo documentation of the whale and an interesting observation. Here I quote the field notes that Dr. Torres wrote from this sighting: “September 6th, 2019. Sighting 9: Scattered whales feeding and/or traveling across area to north of Cape Foulweather. One whale had recently chopped fluke; tried to re-find to get better photos but could not (looking at photos now, this whale is clearly entangled in line!). Ceiling too low for UAS [drone flight].” (Picture 4.B).

Progressive photos of Roller Skate’s fluke. In 2017, her fluke was healthy (A). In 2019, we observed her fluke with a line entanglement and “chewed up” (B). In 2020, we have observed the resulting wound and healing process (C). Photographs captured under NOAA/NMFS permit #21678.

Roller skate’s story is an example of how essential is to keep an ID catalog. After a close-up examination of the 2019 picture, we can clearly see a rope entanglement (Picture 4.B). Photos from previous years show how beautiful and healthy her fluke was before this event (Picture 4.A). This event is heart breaking to witness, but this whale could be considered lucky because she was able to shed the gear and survive this entanglement, at least in the short term. Additionally, we can learn from Roller Skate’s misfortune to help us understand what the consequences of such an injury (stressor) may be on the physiology of a whale. We have been eager to collect a fecal sample from Roller Skate to analyze how her hormone levels compare to non-injured whales. Fortunately, we got lucky a couple weeks ago and collected this sample, so now we need to get in the lab and analyze the samples. But more questions remain: Will this injury impact her ability to reproduce? If so, for how long? And at a larger scale, what are the population consequences of such events? If we can understand the magnitude of lethal and sublethal human caused impacts on individual whales and their populations from events such as entanglements, we can develop better methods to mitigate and limit such hazards for whales in their environments.

As I promised, there is also some good news to share. A very well-known PCFG whale, almost a celebrity I dear to say, is “Scarback”, or as we like to call her “Scarlett”. Scarlett is a female known since 1996, making her at least 24 years old, and she also has a very bad injury of unknown origin. Scarlett has a terrible scar on her back that is theorized to have been caused by an explosive harpoon, or maybe a bad ship-strike (Picture 5), but we really do not know. However, we do know she survived this injury and this year she brought a new calf into the population (Picture 6). This is the second calf we have documented from Scarlett, with her previous calf sighted during the 2016 field season and we call it “Brown”. Scarlett is an example of how resilient these amazing giants can be; however, it is likely that while she was recovering from this injury, she was unable to reproduce. How many calves from Scarlett did the PCFG population “lose” due to such a tragedy? We can´t know, but we are learning, and her story will also help us understand whale physiology as we will analyze her fecal hormones and body condition during pregnancy, lactation, and resting phases.

The scar on Scarlett’s back. This significant wound has an unknown origin. Photograph captured under NOAA/NMFS permit #21678.

Scarlett is a survivor. We need to recognize that we are sharing the ocean with different forms of life. We need to acknowledge their existence and understand how our use of the oceans is affecting them, and, more importantly, work toward improving their conditions. I hope that with our research we highlight and communicate how amazing are these animals, and how important are they for marine ecosystems. And ultimately, I hope our work helps minimize the impacts that affect other forms of ocean life that coexist with us, both above and below the surface.

The well-known gray whale “Scarlett” surfaces under her calf while it checks out the GEMM Lab field team. Photo by Alejandro Fernandez Ajo taken under NOAA/NMFS permit #21678.

References:

Burgess, E., Hunt, K. E., Kraus, S. D. and Rolland, R. M. (2016). Get the most out of blow hormones: validation of sampling materials, field storage and extraction techniques for whale respiratory vapor samples. Conservation Physiology, 4, cow024.

Burgess, E. A., Hunt, K. E., Kraus, S. D. and Rolland, R. M. (2018). Quantifying hormones in exhaled breath for physiological assessment of large whales at sea. Scientific Reports, 8, 10031.

Corkeron, P. J., Rolland, R. M., Hunt, K. E. and Kraus, S. D. (2017). A right whale PooTree: Fecal hormones and classification trees identify reproductive states in North Atlantic right whales (Eubalaena glacialis). Conservation Physiology, 5, cox006. DOI: 10.1093/conphys/cox006.

Hunt, K., Lysiak, N., Moore, M. and Rolland, R. (2017). Multi-year longitudinal profiles of cortisol and corticosterone recovered from baleen of North Atlantic right whales (Eubalaena glacialis). General and Comparative Endocrinology, 254, 50-59. DOI: 10.1016/j.ygcen.2017.09.009.

Hunt, K., Lysiak, N. S. J., Matthews, C. J. D., et al. (2018). Multi-year patterns in testosterone, cortisol and corticosterone in baleen from adult males of three whale species. Conservation Physiology, 6, coy049. DOI: 10.1093/conphys/coy049.

Hunt, K. E., Rolland, R. M., Kraus, S. D. and Wasser, S. K. (2006). Analysis of fecal glucocorticoids in the North Atlantic Right Whale (Eubalaena glacialis). General and Comparative Endocrinology, 148, 260-272.

Lysiak, N., Trumble, S., Knowlton, A. and Moore, M. (2018). Characterizing the duration and severity of fishing gear entanglement on a North Atlantic right whale (Eubalaena glacialis) using stable isotopes, steroid and thyroid hormones in baleen. Frontiers in Marine Science. DOI: 10.3389/fmars.2018.00168.

Rolland, R. M., Hunt, K. E., Kraus, S. D. and Wasser, S. K. (2005). Assessing reproductive status of right whales (Eubalaena glacialis) using fecal hormone metabolites. General and Comparative Endocrinology, 142, 308-317.

Dual cameras provide bigger picture

By Hunter Warick, Research Technician, Geospatial Ecology of Marine Megafauna Lab, Marine Mammal Institute

When monitoring the health of a capital breeding species, such as whales that store energy to support reproduction costs, it is important to understand what processes and factors drive the status of their body condition. Information gained will allow for better insight into their cost of reproduction and overall life history strategies.

For the past four years the GEMM Lab has utilized the perspective that Unoccupied Aerial Systems (UAS; or ‘drones’) provide for observations of marine mammals. This aerial perspective has documented gray whale behavior such as jaw snapping, drooling mud, and headstands, all of which shows or suggest foraging (Torres et al. 2018). However, UAS is limited to a bird’s eye view, allowing us to see WHAT whales are doing, but limited information about the reasons WHY. To overcome this hurdle, Leigh Torres and team have equipped their marine mammal research utility belts with the use of GoPro cameras. They developed a technique known as the “GoPro drop” where a GoPro camera mounted to a weighted pole is lowered off the side of the research vessel in waters < 20 m deep via a line to record video data. This technique allows the team to obtain fine-scale habitat and prey variation information, like what the whale experiences. Along with the context provided by the UAS, this dual camera perspective allows for deeper insight into gray whale foraging strategies and efficiency. Torres’s GoPro data analysis protocol examines kelp density, kelp health, benthic substrate, rock fish density, and mysid density. These characteristics are graded along a scale (Figure 1), allowing for relative comparisons of habitat and prey availability between where whales spend time and forage. These GoPro drops will also help create a fine-scale benthic habitat map of the Newport field area. So, why are these data on gray whale habitat and prey important to understand?

Figure 1. The top row shows varying degrees of mysid density (low to high, left to right). Middle row illustrates different types of substrate you might encounter (reef, sandy, boulders; left to right). Bottom row shows the different levels of kelp health (poor, medium, good).

The foraging grounds are the first step in the life history domino chain reaction for many rorqual whales; if this step doesn’t go off cleanly then everything else fails to fall into place. Gray whales partake on a 15,000-20,000 km (round trip) migration, which is the longest of any known mammal (Swartz 1986). During this migration, whales spend around three months fasting in their breeding grounds (Highsmith & Coyle 1992), living only off the energy stores that they accumulated in their feeding grounds (Næss et al. 1998). These extreme conditions of existence for gray whales drive the need to be a successful forager and is why it is so crucial for them to forage in high prey density areas (Newell, C. 2009).

Mysids are a critical part of the gray whale diet in Oregon waters (Newell, C. 2009; Sullivan, F. 2017) and mysids have strong predator-prey relationships with both top-down and bottom-up control (Dunham & Duffus 2001; Newell & Cowles 2006). This unique tie illustrates the great dependency that gray whales have on mysids, further showing the benefit to looking at the density of mysids where gray whales are seen foraging. The quality of mysids may also be as important as quantity; with higher water temperatures resulting in lower lipid content in mysids (Mauchline 1980), suggesting density might not be the only factor for determining efficient whale foraging. The overall goal of gray whales on their foraging grounds is to get as fat as possible in order to reproduce as often as possible. But, this isn’t always as easy as it sounds. Gray whales typically have a two-year breeding interval but can be anywhere from 1-4 years (Blokhin 1984). The longer time it takes to build up adequate energy stores to support reproduction costs, the longer it will take to breed successfully. Building back up these energy stores can prove to be difficult, especially for lactating females (Figure 2).

Figure 2. Comparison of body condition between a lactating female gray whale on the feeding grounds in Newport, Oregon, 2020 (GEMM Lab, OSU; NOAA/NMFS permit # 21678) and a pregnant female gray whale on the breeding grounds in San Ignacio Lagoon, 2019 (provided by Laguna San Ignacio Ecosystem Program). Photographer Hunter Warick. Note the very different body shapes: thin lactating female relative to the rotund pregnant female.

Being able to track the health and behavior of gray whales on an individual level, including comparisons between variation in body condition, foraging behavior, and fine scale information on benthic communities gained through the use of GoPros, can provide a better understanding of the driving factors and impacts on their health and population trends (Figure 3).


Figure 3. A compilation of video clips captured by the GEMM Lab during their research on gray whale ecology and physiology off Newport, Oregon using Unoccupied Aerial Systems (UAS, or “drones”) and GoPro cameras. UAS are used to observe gray whale behavior and conduct photogrammetry assessment of body condition. GoPro camera drops assess the benthic habitat and prey density across the study region, with a couple chance encounters of whales. Research is conducted under NOAA/NMFS permit # 21678.

Makah Gray Whale Hunt Waiver – a long-time coming, but still premature?

By Lisa Hildebrand, MSc student, OSU Department of Fisheries & Wildlife, Marine Mammal Institute, Geospatial Ecology of Marine Megafauna Lab

Archaeological site of Ozette Village. Source: Makah Museum.

The Makah, an indigenous people of the Pacific Northwest Coast living in Washington State, have a long history with whaling. Deposits from a mudslide in the village of Ozette suggest that whaling may date back 2,000 years as archaeologists uncovered humpback and gray whale bones and barbs from harpoons (Kirk 1986). However, the history of Makah whaling is also quite recent. On January 29 of this year, the National Marine Fisheries Service (NMFS; informally known as NOAA Fisheries) announced a 45-day public comment period regarding a NMFS proposed waiver on the Marine Mammal Protection Act’s (MMPA) moratorium on the take of marine mammals to allow the Makah to take a limited number of eastern North Pacific gray whales (ENP). To understand how the process reached this point, we first must go back to 1855.

1855 marks the year in which the U.S. government and the Makah entered into the Treaty of Neah Bay (in Washington state). The Makah ceded thousands of acres of land to the U.S. government, and in return reserved their right to whale. Following the treaty, the Makah hunt of gray whales continued until the 1920s. At this point, commercial hunting had greatly reduced the ENP population, so much so that the Makah voluntarily ceased their whaling. The next seven decades brought about the formation of the International Whaling Commission (IWC), the enactment of the Whaling Convention Act, the listing of gray whales as endangered under the U.S. Endangered Species Act, and the enactment of the MMPA. For gray whales, these national and international measures were hugely successful, leading to the removal of the ENP from the Federal List of Endangered Wildlife in 1994 when it was determined that the population had recovered to near its estimated original population size.

One year later on May 5, 1995 (just one month after I was born!), the Makah asked the U.S. Department of Commerce to represent its interest to obtain a quota for gray whales from the IWC in order to resume their treaty right for ceremonial and subsistence harvest of the ENP. The U.S. government pursued this request at the next IWC meeting, and subsequently NMFS issued a final Environmental Assessment that found no significant impact to the ENP population if the hunt recommenced. The IWC set a catch limit and NMFS granted the Makah a quota in 1998. In 1999 the Makah hunted, struck and landed an ENP gray whale.

“Makahs cutting up whale, Neah Bay, ca. 1930. Photo by Asahel Curtis, Courtesy UW Special Collections (CUR767)”. Source and caption: History Link.

I will not go into detail about what happened between 1999 and now because frankly, a lot happened, particularly a lot of legal events including summary judgements, appeals, and a lot of other legal jargon that I do not quite understand. If you want to know the specifics of what happened in those two decades, I suggest you look at NMFS’ chronology of the Makah Tribal Whale Hunt. In short, cases brought against NMFS argued that they did not take a “hard [enough] look” at the National Environmental Policy Act when deciding that the Makah could resume the hunt. Consequently, the hunt was put on hold. Yet, in 2005 NMFS received a waiver request from the Makah on the MMPA’s take moratorium and NMFS published a notice of intent to review this request. A lot more happened between that event and now, including on January 29 of this year when NMFS announced the availability of transcripts from the Administrative Law Judge’s (ALJ) hearing (which happened from November 14-21, 2019) on the proposed regulations and waiver to allow the Makah to resume hunting the ENP. We are currently in the middle of the aforementioned 45-day public comment period on the formal rulemaking record. 

It has been 15 years since the Makah requested the waiver and while the decision has not yet been reached, we are likely nearing the end of this long process. This blog has turned into somewhat of a history lesson (not really my intention) but I feel it is important to understand the lengthy and complex history associated with the decision that is probably going to happen sometime this year. My actual intent for this blog is to ruminate on a few questions, some of which remain unanswered in my opinion, that are large and broad, and important to consider. Some of these questions point out gaps in our ecological knowledge regarding gray whales that I believe should be addressed for a truly informed decision to be made on NMFS’ proposed waiver now or anytime in the near future. 

1. Should the Pacific Coast Feeding Group (PCFG) of gray whales be recognized as its own stock?

Currently, the PCFG are considered a part of the ENP stock. This decision was published following a workshop held by a NMFS task force (Weller et al. 2013). The report concluded that based on photo-identification, genetics, tagging, and other data, there was a substantial level of uncertainty in the strength of the evidence to support the independence of the PCFG from the ENP. Nevertheless, mitochondrial genetic data have indicated a differentiation between the PCFG and the ENP, and the exchange rate between the two groups may be small enough for the two to be considered demographically independent (Frasier et al. 2011). Based on all currently available data, it seems that matrilineal fidelity plays a role in creating population structure within and between the PCFG and the ENP, however there has not been any evidence to suggest that whales from one feeding area (i.e. the PCFG range) are reproductively isolated from whales that utilize other feeding areas (i.e. the Arctic ENP feeding grounds) (Lang et al. 2011). Several PCFG researchers do argue that there needs to be recognition of the PCFG as an independent stock. It is clear that more research, especially efforts to link genetic and photo-identification data within and between groups, is required.

ENP gray whales foraging off the coast of Alaska on their main foraging grounds in the Bering Sea. Photo taken by ASAMM/AFSC. Funded by BOEM IAA No. M11PG00033. Source: NMFS.

2. Is emigration/immigration driving PCFG population growth, or is it births/deaths?

It is unclear whether the current PCFG population growth is a consequence of births and deaths that occur within the group (internal dynamics) or whether it is due to immigration and emigration (external dynamics). Likely, it is a combination of the two, however which of the two has more of an effect or is more prevalent? This question is important to answer because if population growth is driven more by external dynamics, then potential losses to the PCFG population due to the Makah hunt may not be as detrimental to the group as a whole. However, if internal dynamics play a bigger role, then the loss of just a few females could have long-term ramifications for the PCFG (Schubert 2019). NMFS has taken precautions to try and avoid such effects. In their proposed waiver, of the cumulative limit of 16 strikes of PCFG whales over the 10-year waiver period, no more than 8 of the strikes may be PCFG females (Yates 2019a). While a great step, it still begs the question how the loss of 8 females, admittedly over a rather long period of time, may affect population dynamics since we do not know what ultimately drives recruitment. Especially when taken together with potential non-lethal effects on whales (further discussed in question 5 below).

“Scarlet” is a PCFG female who has had multiple calves in the decades that researchers have seen her in the PCFG range. Image captured under NOAA/NMFS permit #21678. Source: L Hildebrand.

3. How important are individual patterns within the PCFG, and how might the loss of these individuals affect the population? 

The hunt will be restricted to the Makah Usual & Accustomed fishing area (U&A), which is off the Washington coast. It has been shown that site fidelity among PCFG individuals is strong. In fact, based on the 143 PCFG gray whales observed in nine or more years from 1996 to 2015, 94.4% were seen in at least one of nine different PCFG regions during six or more of the years they were seen (Calambokidis et al. 2017). While high site-fidelity seems to be common for some PCFG individuals in certain regions, interestingly, an analysis of sighting histories of all individuals that utilized the Makah U&A from 1985-2011 revealed that most PCFG whales do not have strong site fidelity to the Makah U&A (Scordino et al. 2017). Only about 20% of the whales were observed in six or more years of the total 26 years of data analyzed. Since high individual site fidelity does not appear to be strong in this area, perhaps a loss of genetic diversity, cultural knowledge, and behavioral individualism is not of great concern.

“Buttons” seems to have a preference for the southern Oregon coast as in the last 5 years the GEMM Lab has conducted research, he has only been sighted in 1 year in Newport but in all 5 years in Port Orford. However, perhaps such preferences are not common among all PCFG whales. Source: F. Sullivan.

4. How has the current UME affected the situation?

The ENP has experienced two Unusual Mortality Events (UMEs) in the past 20 years; one from 1999-2000 and the second began in May 2019. Many questions arise when thinking about the Makah hunt in light of the UME. 

  • What impacts will the current UME have on ENP and PCFG birth rates in subsequent years? 
  • Could the UME lead to shifts in feeding behavior of ENP whales and result in greater use of PCFG range by more individuals?
  • What caused the UME? Shifting prey availability and a changing climate? Or has the ENP reached carrying capacity? 
  • Will UMEs become more frequent in the future with continued warming of the Arctic? 
  • What is the added impact of such periodic UMEs on population trends?
“A gray whale found dead off Point Reyes National Seashore in northern California [during the 2019 UME]. Photo by M. Flannery, California Academy of Sciences.” Source and caption: NMFS.

A key assumption of the model developed by NMFS (Moore 2019) to forecast PCFG population size for the period 2016-2028, is that the population processes underlying the data from 2002-2015 (population size estimates developed by Calambokidis et al. 2017) will be the same during the forecasted period. In other words, it is assuming that PCFG gray whales will experience similar environmental conditions (with similar variation) during the next decade as the previous one, and that there will be no catastrophic events that could drastically affect population dynamics. The UME that is still ongoing could arguably affect population dynamics enough such that they are drastically different to effects on the population dynamics during the previous decade. The cause of  the 1999/2000 UME remains undetermined and the results of the investigation of the current UME will possibly not be available for several years (Yates 2019b). Even though the ENP did rebound following the 1999/2000 UME and the abundance of the PCFG increased during and subsequent to that UME, much has changed in the 20 years since then. Increased noise due to increased vessel traffic and other anthropogenic activities (seismic surveys, pile driving, construction to name a few) as well as increased coastal recreational and commercial fishing, have all contributed to a very different oceanscape than the ENP and PCFG encountered 20 years ago. Furthermore, the climate has changed considerably since then too, which likely has caused changes in the spatial distribution of habitat and quantity, quality, and predictability of prey. All of these factors make it difficult to predict what impact the UME will have now. If such events were to become more frequent in the future or the impacts of such events are greater than anticipated, then the PCFG population forecasts will not have accounted for this change. 

5. What impacts will the hunt and associated training exercises have on energy and stress levels of whales?

The proposed waiver would allow hunts to occur in the following manner: in even-years, the hunting period is from December 1 of an odd-numbered year through May 31 of the following even-numbered year. While in odd-years, the hunt is limited from July to October.

In the even-years, the hunt coincides with the northbound migration toward the foraging grounds for ENP whales and with the arrival of PCFG whales to their foraging grounds near the Makah U&A. During the northbound migration, gray whales are at their most nutritionally stressed state as they have been fasting for several months. They are therefore most vulnerable to energy losses due to disturbance at this point (Villegas-Amtmann 2019). Attempted strikes and training exercises would certainly cause some level of disturbance and stress to the whales. Furthermore, the timing of even-year hunts, means that hunters would likely encounter pregnant females, as they are the first to arrive at foraging grounds. A loss of just ~4% of a pregnant female’s energy budget could cause them to abort the fetus or not produce a calf that year (Villegas-Amtmann 2019).

In odd-years, the Makah hunt will most certainly target PCFG whales as the Makah U&A forms one of the nine PCFG regions where PCFG individuals will be feeding during those months. However, NMFS’ waiver limits the number of strikes during odd-years to 2 (Yates 2019a), which certainly protects the PCFG population.

Stress is a difficult response to quantify in baleen whales and research on stress through hormone analysis is still relatively novel. It is unlikely that a single boat training approach of a gray whale will have an adverse effect on the individual. However, a whale is never just experiencing one disturbance at a time. There are typically many confounding factors that influence a whale’s state. In an ideal world, we would know what all of these factors are and how to recognize these effects. Yet, this is virtually impossible. Therefore, while precautions will be taken to try to minimize harm and stress to the gray whales, there may very well still be unanticipated impacts that we cannot anticipate. 

Gray whale fluke. Image captured under NOAA/NMFS permit #21678. Photo: L Hildebrand.

Final thoughts

Many unknowns still remain about the ENP and PCFG gray whale populations. During the ALJ hearing, both sides tried to deal with these unknowns. After reading testimony from both sides, it is clear to me that some of the unknowns still have not been reconciled. Ultimately, a lot of the questions circle back to the first one I posed above: Are the PCFG an independent stock? If there is independent population structure, then the proposed waiver put forth by NMFS would likely change. While NMFS has certainly taken the PCFG into account during the declarations of several experts at the ALJ hearing and has aired on the side of caution, the fact that the PCFG is considered part of the ENP might underestimate the impact that a resumption of the Makah hunt may have on the PCFG. As you can see, there are still many questions that should be addressed to make fully informed decisions on such an important ruling. While this research may take several years to obtain results, the data are within reach through synthesis and collaboration that will fill these critical knowledge gaps. 

Literature cited

Calambokidis, J. C., J. Laake, and A. Pérez. 2017. Updated analysis of abundance and population structure of seasonal gray whales in the Pacific Northwest, 1996-2015. International Whaling Commission SC/A17/GW/05.

Frasier, T. R., S. M. Koroscil, B. N. White, and J. D. Darling. 2011. Assessment of population substructure in relation to summer feeding ground use in eastern North Pacific gray whale. Endangered Species Research 14:39-48.

Kirk, Ruth. 1986. Tradition and change on the Northwest Coast: the Makah, Nuu-chah-nulth, southern Kwakiutl and Nuxalk. University of Washington Press, Seattle.

Lang, A. R., D. W. Weller, R. LeDuc, A. M. Burdin, V. L. Pease, D. Litovka, V. Burkanov, and R. L. Brownell, Jr. 2011. Genetic analysis of stock structure and movements of gray whales in the eastern and western North Pacific. SC/63/BRG10.

Moore, J. E. 2019. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Schubert, D. J. 2019. Rebuttal testimony in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Scordino, J. J., M. Gosho, P. J. Gearin, A. Akmajian, J. Calambokidis, and N. Wright. 2017. Individual gray whale use of coastal waters off northwest Washington during the feeding season 1984-2011: Implications for management. Journal of Cetacean Research and Management 16:57-69.

Villegas-Amtmann, S. 2019. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001.

Weller, D. W., S. Bettridge, R. L. Brownell, Jr., J. L. Laake, J. E. Moore, P. E. Rosel, B. L. Taylor, and P. R. Wade. 2013. Report of the National Marine Fisheries Service Gray Whale Stock Identification Workshop. NOAA-TM-NMFS-SWFSC-507. 

Yates, C. 2019a. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Yates, C. 2019b. Fifth declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.