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

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

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

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

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

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

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

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

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

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


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

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

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

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

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

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

Keeping up with the HALO project: Recovering Rockhopper acoustic recording units and eavesdropping on Northern right whale dolphins

Marissa Garcia, PhD Student, Cornell University, Department of Natural Resources and the Environment, K. Lisa Yang Center for Conservation Bioacoustics

It was a June morning on the Pacific Ocean, and the R/V Pacific Storm had come to a halt on its journey back to shore. The night before, the Holistic Assessment of Living marine resources off Oregon (HALO) project team had disembarked from Newport and began the long transit to NH 65, a site 65 nautical miles offshore along the Newport Hydrographic line (NH line). Ever since the 1960s, researchers have been conducting oceanographic studies along the NH line; the HALO project seeks to add the biological dimension to these historical data collections.

We were on a mission to recover our first set of Rockhoppers that we had deployed in October 2021, just nine months earlier. The Rockhopper is an underwater passive acoustic recording unit developed by K. Lisa Yang Center for Conservation Bioacoustics at Cornell University. Earlier versions of underwater recorders were optimized to record baleen whales. By contrast, the Rockhopper is designed to record both baleen whales and dolphins on longer and deeper deployments, making it apt for research endeavors such as the HALO project. Three units, deployed at NH 25, 45, and 65, continuously recorded the soundscape of the Oregon waters for six months. In June, we were headed out to sea to recover these three units, collect the acoustic data, and deploy three new units.

Figure 1: The HALO project routinely surveys the trackline spanning between NH 25 and NH 65 on the NH line. Credit: Leigh Torres.

With the ship paused, our first task was to recover the Rockhopper we had deployed at NH 65. This Rockhopper deployment at NH 65 was our deepest successful deployment to date, moored at nearly 3,000 m.

So, how does one recover an underwater recording unit that is nearly 3,000 m below the surface? When the Rockhopper was deployed, it was anchored to the seafloor with a 60 kg cast iron anchor. It seems improbable that an underwater recording unit — anchored by such heavy weights — can eventually rise to the surface, but this capability is made possible through a piece of attached equipment called the acoustic release. By sending a signal of a numbered code from a box on the boat deck through the water column to the Rockhopper, the bottom of the acoustic release will begin to spin and detach from the weights. The weights are then left on the seafloor, as the Rockhopper slowly rises to the surface, now unhindered by the weights. Since these weights are composed of iron, they will naturally erode, without additional pollution contributed to the ecosystem. At NH 65, it took approximately an hour for the Rockhopper to reach the surface.

Figure 2: A diagram of the Rockhopper mooring. Of particular importance to this blog post is the acoustic release (Edgtech PORT MFE release) and the 60 kg anchor (Source: Klinck et al., 2020).

The next challenge is finding the Rockhopper bobbing amongst the waves in the vast ocean — much like searching for a needle in a haystack. The color of the Rockhopper helps aid in this quest. It’s imperative anyone out on the boat deck wears a life jacket; if someone goes overboard while wearing a life-jacket, on-board passengers can more easily spot a bright orange spot in an otherwise blue-green ocean with white caps. The design of the Rockhopper functions similarly; the unit is contained in a bright orange hard hat, helping researchers on-board to more easily spot the device, especially in an ocean often characterized by high sea state.

We also use a Yagi antenna to listen for the VHF (Very High Frequency) signal of the recovery gear, a signal the Rockhopper emits once it’s surfaced above the waterline. Pointing the antenna toward the ocean, we can detect the signal, which will become stronger when we point antenna in the direction of the Rockhopper; once we hear that strong signal, we can recommend to the boat captain to start moving the vessel in that direction.

Figure 3: Derek Jaskula, a member of the field operations team at the K. Lisa Yang Center for Conservation Bioacoustics, points the Yagi antenna to detect the signal from the surfaced Rockhopper. Credit: Marissa Garcia.

At that point, all eyes are on the water, binoculars scanning the horizon for the orange. All ears are eager for the exciting news: “I see the Rockhopper!”

Once that announcement is made, the vessel carefully inches toward the Rockhopper until it is just next to the vessel’s side. Using a hook, the Rockhopper is pulled upward and back onto the deck.

What we weren’t expecting, however, during this recovery was to have our boat surrounded by two dolphin species: Pacific white-sided dolphins (Lagenorhynchus obliquidens) and Northern right whale dolphins (Lissodelphis borealis).

One HALO team member shouted, “I see Northern right whale dolphins!”

Charged with excitement, I quickly climbed up the crow’s nest to get a birds-eye look at the ocean bubbling around us with surfacing dolphins. Surely enough, I spotted the characteristic stripe of the Pacific-white sided dolphins zooming beneath the surface, in streaks of white. But what I was even more eager to see were the Northern right whale dolphins, flipping themselves out of the water, unveiling their bright white undersides. Because they lack dorsal fins, we on-board colloquially refer to Northern right whale dolphins as “sea slugs” to describe their appearance as they surface.

Figure 4: The Northern right whale dolphin (Lissodelphis borealis) surfaces during a HALO cruise. Source: HALO Project Team Member. Permit: NOAA/NMFS permit #21678.

In my analysis of the HALO project data for my PhD, I am interested in using acoustics to describe how the distribution of dolphins and toothed whales in Oregon waters varies across space and time. One species I am especially fascinated to study in-depth is the Northern right whale dolphin. To my knowledge, only three papers to date have attempted to describe their acoustics — two of which were published in the 1970s, and the most recent of which was published fifteen years ago (Fish & Turl, 1976; Leatherwood & Walker, 1979; Rankin et al., 2007).

Leatherwood & Walker (1979) proposed that Northern right whale dolphins produced two categories of whistles: a high frequency whistle that turned into burst-pulse vocalizations, and low frequency whistles. However, Rankin et al. (2007) proposed that Northern right whale dolphins may not actually produce whistles, based on two lines of evidence. First, Rankin et al. (2007) combined visual and acoustic survey, and all vocalizations recorded were localized via beamforming methods to verify that recorded vocalizations were produced by the visually observed dolphins. The visual surveying component is key to validating the vocalizations of the species, which also hints that the HALO project’s multi-surveying approach (acoustic and visual) could help arrive at similar results. Second, the Rankin et al. (2007) explored the taxonomy of the Northern right whale dolphin to verify which vocalizations the species is likely to produce based on the vocal repertoire of its close relatives. The right whale dolphin is closely related to dolphins in the genus Lagenorhynchus — which includes white-sided dolphins — and Cephalorhynchus — which includes Hector’s dolphin. The vocal repertoire of these relatives don’t produce whistles, and instead predominantly produced pulsed sounds or clicks (Dawson, 1991; Herman & Tavolga, 1980). Northern right whale dolphins primarily produce echolocation clicks trains and burst-pulses. Although Rankin et al. (2007) claims that the Northern right whale dolphin does not produce whistles, stereotyped burst-pulse series may be unique to individuals, just as dolphin species use stereotyped signature whistles, or they may be relationally shared just as discrete calls of killer whales are.

Figure 5: The Northern right whale dolphin (Lissodelphis borealis) produces burst-pulses. There exists variation in series of burst-pulses. The units marked by (a) and (b) ultimately get replaced by the unit marked by (c). (Source: Rankin et al., 2007).

We have just finished processing the first round of acoustic data for the HALO project, and it is ready now for analysis. Already previewing an hour of data on the Rockhopper by NH 25, we identified potential Northern right whale dolphin recordings . So far, we have only visually observed Northern right whale dolphins nearby Rockhopper units placed at sites NH 65 and NH 45, so it was surprising to acoustically detect this species on the most inshore unit at NH 25. I look forward to demystifying the mystery of Northern right whale dolphin vocalizations as our research on the HALO project continues!

Figure 6: Potential Northern right whale dolphin vocalizations recorded at the Rockhopper deployed at NH 25.

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Dawson, S. (1991). Clicks and Communication: The Behavioural and Social Contexts of Hector’s Dolphin Vocalizations. Ethology, 88(4), 265–276.

Fish, J. F. & Turl, C. W. (1976). Acoustic Source Levels of Four Species of Small Whales.

Herman, L. M., and Tavolga, W. N. (1980). “The communication systems of cetaceans,” in Cetacean behavior: Mechanisms and functions, edited by L. M. Herman (Wiley, New York), 149–209.

Klinck, H., Winiarski, D., Mack, R. C., Tessaglia-Hymes, C. T., Ponirakis, D. W., Dugan, P. J., Jones, C., & Matsumoto, H. (2020). The Rockhopper: a compact and extensible marine autonomous passive acoustic recording system. Global Oceans 2020: Singapore – U.S. Gulf Coast, 1–7.

Leatherwood, S., and Walker, W. A. (1979). “The northern right whale dolphin Lissodelphis borealis Peale in the eastern North Pacific,” in Behavior of marine animals, Vol. 3: Cetaceans, edited by H. E. Winn and B. L. Olla (Plenum, New York), 85–141.

Rankin, S., Oswald, J., Barlow, J., & Lammers, M. (2007). Patterned burst-pulse vocalizations of the northern right whale dolphin, Lissodelphis borealis. The Journal of the Acoustical Society of America, 121(2), 1213–1218.

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

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


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

Where Gray Whales Are and Why It Matters

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

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

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

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


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

Table 1. Hypotheses and Expected Results.

A Hundred and One Data Visualizations

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

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

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

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


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


Benoit-Bird, K. J., & McManus, M. A. (2012). Bottom-up regulation of a pelagic community through spatial aggregations. Biology Letters8(5), 813–816.

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

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

Harbor porpoise and gray whale distribution over three decades: introducing the EMERALD project

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

Throughout the world, humans rely on coastal regions for shipping and commerce, fisheries, industrial development, and increasingly for the development of marine renewable energy such as wind and wave energy [1]. Nearshore environments, including the coastal waters of the Northern California Current (NCC), are therefore coupled social-ecological systems, at the intersection of human and biological productivity [2].

The NCC supports a diverse food web of ecologically and commercially important species [3]. The nearshore region of the NCC is further shaped by a rich mosaic of complex features including rocky reefs, kelp forests, and sloping sandy bottom substrate [4], creating habitat for numerous species of conservation interest, including invertebrates, fish, seabirds, and marine mammals [5]. Despite its importance, this realm poses significant challenges for vessel-based data collection, and therefore it remains relatively poorly monitored and understood.

The view from Cape Foulweather, showing the complex mosaic of nearshore habitat features. Photo: D. Barlow.

I am excited to introduce a new project focused on these important nearshore waters, in which we will be Examining Marine mammal Ecology through Region-wide Assessment of Long-term Data (EMERALD). Since 1992, standardized surveys have been conducted between San Francisco Bay, CA, and the Columbia River, OR, to monitor the abundance of marbled murrelets, a seabird of conservation concern. Each spring and summer, researchers have simultaneously been diligently documenting the locations of harbor porpoise and gray whale sightings—two iconic marine mammal species that rely on the nearshore waters of the NCC. This rich and extensive record is rare for marine mammal data, particularly in the challenging, turbulent nearshore environment. Furthermore, harbor porpoises are cryptic, making visual sampling particularly challenging, and gray whales can be sparsely distributed, yielding low sample sizes in the absence of long-term data collection.

Left: The survey team collecting data; Right: Marbled murrelet floating on the water.

For the EMERALD project, we will investigate spatial and temporal distribution patterns of harbor porpoises and gray whales in relation to fluctuations in key environmental drivers. The primary goals of the project are to (1) Identify persistent hotspots in harbor porpoise and gray whale sightings over time, and (2) Examine the environmental drivers of sighting hotspots through spatial and temporal analyses.

A harbor porpoise surfacing off the central Oregon coast. Photo: L. Torres.

From a first look at the data, we are already excited by some emerging patterns. In total, the dataset contains sightings of 6,763 harbor porpoise (mean 233 per year) and 530 gray whales (mean 18 per year). Preliminary data exploration reveals that harbor porpoise sightings increased in 2011-2012, predominantly between Cape Blanco, OR, and Cape Mendocino, CA. Gray whale sightings appear to follow an oscillating, cyclical pattern with peaks approximately every three years, with notable disruption of this pattern during the marine heatwave of 2014-2015. What are the drivers of sighting hotspots and spatial and temporal fluctuations in sighting rates? Time—and a quantitative analytical approach involving density estimation, timeseries analysis, and species distribution modeling—will tell.

A gray whale forages in kelp forest habitat over a nearshore rocky reef. Photo: T. Chandler.

I recently completed my PhD on the ecology and distribution of blue whales in New Zealand (for more information, see the OBSIDIAN project). Now, I am excited to apply the spatial analysis skills have been honing to a new study system and two new study species as I take on a new role in the GEMM Lab as a Postdoctoral Scholar. The EMERALD project will turn my focus to the nearshore waters close to home that I have grown to love over the past six years as a resident of coastal Oregon. The surveys I will be working with began before I was born, and I am truly fortunate to inherit such a rich dataset—a rare treat for a marine mammal biologist, and an exciting prospect for a statistical ecologist.

Dawn and Quin the dog, enjoying views of Oregon’s complex and important nearshore waters. Both are thrilled to remain in Oregon for the EMERALD project. Photo: R. Kaplan.

So, stay tuned for our findings as the project unfolds. In the meantime, I want express gratitude to Craig Strong of Crescent Coastal Research who has led the dedicated survey effort for the marbled murrelet monitoring program, without whom none of the data would exist. This project is funded by the Oregon Gray Whale License Plate funds, and we thank the gray whale license plate holders for their support of marine mammal research.

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1.        Jouffray, J.-B., Blasiak, R., Norström, A. V., Österblom, H., and Nyström, M. (2020). The Blue Acceleration: The Trajectory of Human Expansion into the Ocean. One Earth 2, 43–54.

2.        Sjostrom, A.J.C., Ciannelli, L., Conway, F., and Wakefield, W.W. (2021). Gathering local ecological knowledge to augment scientific and management understanding of a living coastal resource: The case of Oregon’s nearshore groundfish trawl fishery. Mar. Policy 131, 104617.

3.        Bograd, S.J., Schroeder, I., Sarkar, N., Qiu, X., Sydeman, W.J., and Schwing, F.B. (2009). Phenology of coastal upwelling in the California Current. Geophys. Res. Lett. 36, 1–5.

4.        Romsos, G., Goldfinger, C., Robison, R., Milstein, R., Chaytor, J., and Wakefield, W. (2007). Development of a regional seafloor surficial geologic habitat map for the continental margins of Oregon and Washington, USA. Mapp. Seafloor Habitat Charact. Geol. Assoc. Canada, Spec. Pap., 219–243.

5.        Oregon Department of Fish and Wildlife (2016). Oregon Nearshore Strategy. Available at: [Accessed January 10, 2022].

Reuniting with some old friends: The 8th GRANITE field season is underway

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

We are almost halfway through June which means summer has arrived! Although, here on the Oregon coast, it does not entirely feel like it. We have been swinging between hot, sunny days and cloudy, foggy, rainy days that are reminiscent of those in spring or even winter. Despite these weather pendulums, the GEMM Lab’s GRANITE project is off to a great start in its 8th field season! The field team has already ventured out onto the Pacific Ocean in our trusty RHIB Ruby on four separate days looking for gray whales and in this blog post, I am going to share what we have seen so far.

The core GRANITE field team before the May 24th “trial run”. From left to right: Leigh Torres, KC Bierlich, Clara Bird, Lisa Hildebrand, Alejandro Fernández Ajó. Source: L. Torres.

PI Leigh, PhD candidate Clara and I headed out for a “trial run” on May 24th. While the intention for the day was to make sure all our gear was running smoothly and we still remembered how to complete the many tasks associated with our field work (boat loading and trailering, drone flying and catching, poop scooping, data download, to name a few), we could not resist surveying our entire study range given the excellent conditions. It was a day that all marine field scientists hope for – low winds (< 5 kt all day) and a 3 ft swell over a long period. Despite surveying between Waldport and Depoe Bay, we only encountered one whale, but it was a whale that put a smile on each of our faces. After “just” 252 days, we reunited with Solé, the star of our GRANITE dataset, with record numbers of fecal samples and drone flights collected. This record is due to what seems to be a strong habitat or foraging tactic preference by Solé to remain in a relatively small spatial area off the Oregon coast for most of the summer, rather than traveling great swaths of the coast in search for food. Honest truth, on May 24th we found her exactly where we expected to find her. While we did not collect a fecal sample from her on that day, we did perform a drone flight, allowing us to collect a critical early feeding season data point on body condition. We hope that Solé has a summer full of mysids on the Oregon coast and that we will be seeing her often, getting rounder each time!

Our superstar whale Solé. Her identifying features are a small white line on her left side (green box) and a white dot in front of her dorsal hump on the right side (red circle). Source: GEMM Lab. Photograph captured under NOAA/NMFS permit #21678

Just a week after this trial day, we had our official start to the field season with back-to-back days on the water. On our first day, postdoc Alejandro, Clara and I were joined by St. Andrews University Research Fellow Enrico Pirotta, who is another member of the GRANITE team. Enrico’s role in the GRANITE project is to implement our long-term, replicate dataset into a framework called Population consequences of disturbance (PCoD; you can read all about it in a previous blog). We were thrilled that Enrico was able to join us on the water to get a sense for the species and system that he has spent the last several months trying to understand and model quantitatively from a computer halfway across the world. Luckily, the whales sure showed up for Enrico, as we saw a total of seven whales, all of which were known individuals to us! Several of the whales were feeding in water about 20 m deep and surfacing quite erratically, making it hard to get photos of them at times. Our on-board fish finder suggested that there was a mid-water column prey layer that was between 5-7 m thick. Given the flat, sandy substrate the whales were in, we predicted that these layers were composed of porcelain crab larvae. Luckily, we were able to confirm our hypothesis immediately by dropping a zooplankton net to collect a sample of many porcelain crab larvae. Porcelain crab larvae have some of the lowest caloric values of the nearshore zooplankton species that gray whales likely feed on (Hildebrand et al. 2021). Yet, the density of larvae in these thick layers probably made them a very profitable meal, which is likely the reason that we saw another five whales the next day feeding on porcelain crab larvae once again.

On our most recent field work day, we only encountered Solé, suggesting that the porcelain crab swarms had dissipated (or had been excessively munched on by gray whales), and many whales went in search for food elsewhere. We have done a number of zooplankton net tows across our study area and while we did collect a good amount of mysid shrimp already, they were all relatively small. My prediction is that once these mysids grow to a more profitable size in a few days or weeks, we will start seeing more whales again.

The GRANITE team from above, waiting & watching for whales, as we will be doing for the rest of the summer! Source: GEMM Lab.

So far we have seen nine unique individuals, flown the drone over eight of them, collected fecal samples from five individuals, conducted 10 zooplankton net tows and seven GoPro drops in just four days of field work! We are certainly off to a strong start and we are excited to continue collecting rock solid GRANITE data this summer to continue our efforts to understand gray whale ecology and physiology.


Literature cited

Hildebrand L, Bernard KS, Torres LGT. 2021. Do gray whales count calories? Comparing energetic values of gray whale prey across two different feeding grounds in the Eastern North Pacific. Frontiers in Marine Science 8. doi: 10.3389/fmars.2021.683634

Yonder Whales and Nearby Prey: A New Look at a Familiar System

Rachel Kaplan1, Dawn Barlow2, Clara Bird3

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

2Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

3PhD Candidate, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

What do peanut butter m&ms, killer whales, affogatos, tired eyes, and puffins all have in common? They were all major features of the recent Northern California Current (NCC) ecosystem survey cruise. 

The science party of the May 2022 Northern California Current ecosystem cruise.

We spent May 6–17 aboard the NOAA vessel Bell M. Shimada in northern California, Oregon, and Washington waters. This fabulously interdisciplinary cruise studies multiple aspects of the NCC ecosystem three times per year, and the GEMM lab has put marine mammal observers aboard since 2018.

This cruise was a bit different than usual for the GEMM lab: we had eyes on both the whales and their prey. While Dawn Barlow and Clara Bird observed from sunrise to sunset to sight and identify whales, Rachel Kaplan collected krill data via an echosounder and samples from net tows in order to learn about the preyscape the whales were experiencing. 

From left, Rachel, Dawn, and Clara after enjoying some beautiful sunset sightings. 

We sailed out of Richmond, California and went north, sampling as far north as La Push, Washington and up to 200 miles offshore. Despite several days of challenging conditions due to wind, rain, fog, and swell, the team conducted a successful marine mammal survey. When poor weather prevented work, we turned to our favorite hobbies of coding and snacking.

Rachel attends “Clara’s Beanbag Coding Academy”.

Cruise highlights included several fin whales, sperm whales, killer whales, foraging gray whales, fluke slapping and breaching humpbacks, and a visit by 60 pacific white-sided dolphins. While being stopped at an oceanographic sampling station typically means that we take a break from observing, having more time to watch the whales around us turned out to be quite fortunate on this cruise. We were able to identify two unidentified whales as sei whales after watching them swim near us while paused on station. 

Marine mammal observation segments (black lines) and the sighting locations of marine mammal species observed during the cruise.

On one of our first survey days we also observed humpbacks surface lunge feeding close to the ship, which provided a valuable opportunity for our team to think about how to best collect concurrent prey and whale data. The opportunity to hone in on this predator-prey relationship presented itself in a new way when Dawn and Clara observed many apparently foraging humpbacks on the edge of Heceta Bank. At the same time, Rachel started observing concurrent prey aggregations on the echosounder. After a quick conversation with the chief scientist and the officers on the bridge, the ship turned around so that we could conduct a net tow in order to get a closer look at what exactly the whales were eating.

Success! Rachel collects krill samples collected in an area of foraging humpback whales.

This cruise captured an interesting moment in time: southerly winds were surprisingly common for this time of year, and the composition of the phytoplankton and zooplankton communities indicated that the seasonal process of upwelling had not yet been initiated. Upwelling brings deep, cold, nutrient-rich waters to the surface, generating a jolt of productivity that brings the ecosystem from winter into spring. It was fascinating to talk to all the other researchers on the ship about what they were seeing, and learn about the ways in which it was different from what they expected to see in May.

Experiencing these different conditions in the Northern California Current has given us a new perspective on an ecosystem that we’ve been observing and studying for years. We’re looking forward to digging into the data and seeing how it can help us understand this ecosystem more deeply, especially during a period of continued climate change.

The total number of each marine mammal species observed during the cruise.

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The pathway to advancing knowledge of rorqual whale distribution off Oregon

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

In September 2020, I was hired as a postdoc in the GEMM Lab and was tasked to conduct the analyses necessary for the OPAL project. This research project has the ambitious, yet essential, goal to fill a knowledge gap hindering whale conservation efforts locally: where and when do whales occur off the Oregon coast? Understanding and predicting whale distribution based on changing environmental conditions is a key strategy to assess and reduce spatial conflicts with human activities, specifically the risk of entanglement in fixed fishing gear.

Starting a new project is always a little daunting. Learning about a new region and new species, in an alien research and conservation context, is a challenge. As I have specialized in data science over the last couple of years, I have been confronted many times with the prospect of working with massive datasets collected by others, from which I was asked to tease apart the biases and the ecological patterns. In fact, I have come to love that part of my job: diving down the data rabbit hole and making my way through it by collaborating with others. Craig Hayslip, faculty research assistant in MMI, was the observer who conducted the majority of the 102 helicopter surveys that were used for this study. During the analysis stage, his help was crucial to understand the data that had been collected and get a better grasp of the field work biases that I would later have to account for in my models. Similarly, it took hours of zoom discussions with Dawn Barlow, the GEMM lab’s latest Dr, to be able to clean and process the 75 days of survey effort conducted at sea, aboard the R/V Shimada and Oceanus.

Once the data is “clean”, then comes the time for modeling. Running hundreds of models, with different statistical approaches, different environmental predictors, different parameters etc. etc. That is when you realize what a blessing it is to work with a supervisor like Leigh Torres, head of the GEMM Lab. As an early career researcher, I really appreciate working with people who help me take a step back and see the bigger picture within which the whole data wrangling work is included. It is so important to have someone help you stay focused on your goals and the ecological questions you are trying to answer, as these may easily get pushed back to the background during the data analysis process.

And here we are today, with the first scientific publication from the OPAL project published, a little more than three years after Leigh and Craig started collecting data onboard the United States Coast Guard helicopters off the coast of Oregon in February 2019. Entitled “Seasonal, annual, and decadal distribution of three rorqual whale species relative to dynamic ocean conditions off Oregon, USA”, our study published in Frontiers in Marine Science presents modern and fine-scale predictions of rorqual whale distribution off Oregon, as well as a description of their phenology and a comparison to whale numbers observed across three decades in the region (Figure 1). This research focuses on three rorqual species sharing some ecological and biological traits, as well as similar conservation status: humpback whales (Megaptera novaeangliae), blue whales (Balaenoptera musculus musculus), and fin whales (Balaenoptera physallus); all of which migrate and feed over the US West coast (see a previous blog to learn more about these species here).

Figure 1: Graphical abstract of our latest paper published in Frontiers in Marine Science.

We demonstrate (1) an increase in rorqual numbers over the last three decades in Oregon waters, (2) differences in timing of migration and habitat preferences between humpback, blue, and fin whales, and (3) predictable relationships of rorqual whale distribution based on dynamic ocean conditions indicative of upwellings and frontal zones. Indeed, these ocean conditions are likely to provide suitable biological conditions triggering increased prey abundance. Three seasonal models covering the months of December-March (winter model), April-July (spring) and August-November (summer-fall) were generated to predict rorqual whale densities over the Oregon continental shelf (in waters up to 1,500 m deep). As a result, maps of whale densities can be produced on a weekly basis at a resolution of 5 km, which is a scale that will facilitate targeted management of human activities in Oregon. In addition, species-specific models were also produced over the period of high occurrence in the region;  that is humpback and blue whales between April and November, and fin whales between August and March. 

As we outline in our concluding remarks, this work is not to be considered an end-point, but rather a stepping stone to improve ecological knowledge and produce operational outputs that can be used effectively by managers and stakeholders to prevent spatial conflict between whales and human activities. As of today, the models of fin and blue whale densities are limited by the small number of observations of these two species over the Oregon continental shelf. Yet, we hope that continued data collection via fruitful research partnerships will allow us to improve the robustness of these species-specific predictions in the future. On the other hand, the rorqual models are considered sufficiently robust to continue into the next phase of the OPAL project that aims to assess overlap between whale distribution and Dungeness crab fishing gear to estimate entanglement risk. 

The curse (or perhaps the beauty?) of species distribution modeling is that it never ends. There are always new data to be added, new statistical approaches to be tested, and new predictions to be made. The OPAL models are no exception to this rule. They are meant to be improved in future years, thanks to continued helicopter and ship-based survey efforts, and to the addition of new environmental variables meant to better predict whale habitat selection. For instance, Rachel Kaplan’s PhD research specifically aims at understanding the distribution of whales in relation to krill. Her results will feed into the more general efforts to model and predict whale distribution to inform management in Oregon.

This first publication therefore paves the way for more exciting and impactful research!

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Derville, S., Barlow, D. R., Hayslip, C. E., and Torres, L. G. (2022). Seasonal, Annual, and Decadal Distribution of Three Rorqual Whale Species Relative to Dynamic Ocean Conditions Off Oregon, USA. Front. Mar. Sci. 9, 1–19. doi:10.3389/fmars.2022.868566.


We gratefully acknowledge the immense contribution of the United State Coast Guard sectors North Bend and Columbia River who facilitated and piloted our helicopter surveys. We would like to also thank NOAA Northwest Fisheries Science Center for the ship time aboard the R/V Bell M. Shimada. We thank the R/V Bell M. Shimada (chief scientists J. Fisher and S. Zeman) and R/V Oceanus crews, as well as the marine mammal observers F. Sullivan, C. Bird and R. Kaplan. We give special recognition and thanks to the late Alexa Kownacki who contributed so much in the field and to our lives. We also thank T. Buell and K. Corbett (ODFW) for their partnership over the OPAL project. We thank G. Green and J. Brueggeman (Minerals Management Service), J. Adams (US Geological Survey), J. Jahncke (Point blue Conservation), S. Benson (NOAA-South West Fisheries Science Center), and L. Ballance (Oregon State University) for sharing validation data. We thank J. Calambokidis (Cascadia Research Collective) for sharing validation data and for logistical support of the project. We thank A. Virgili for sharing advice and custom codes to produce detection functions.

New publication by GEMM Lab reveals sub-population health differences in gray whales 

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

In a previous blog, I discussed the importance of incorporating measurement uncertainty in drone-based photogrammetry, as drones with different sensors, focal length lenses, and altimeters will have varying levels of measurement accuracy. In my last blog, I discussed how to incorporate photogrammetric uncertainty when combining multiple measurements to estimate body condition of baleen whales. In this blog, I will highlight our recent publication in Frontiers in Marine Science ( led by GEMM Lab’s Dr. Leigh TorresClara Bird, and myself that used these methods in a collaborative study using imagery from four different drones to compare gray whale body condition on their breeding and feeding grounds (Torres et al., 2022).

Most Eastern North Pacific (ENP) gray whales migrate to their summer foraging grounds in Alaska and the Arctic, where they target benthic amphipods as prey. A subgroup of gray whales (~230 individuals) called the Pacific Coast Feeding Group (PCFG), instead truncates their migration and forages along the coastal habitats between Northern California and British Columbia, Canada (Fig. 1). Evidence from a recent study lead by GEMM Lab’s Lisa Hildebrand (see this blog) found that the caloric content of prey in the PCFG range is of equal or higher value than the main amphipod prey in the Arctic/sub-Arctic regions (Hildebrand et al., 2021). This implies that greater prey density and/or lower energetic costs of foraging in the Arctic/sub-Arctic may explain the greater number of whales foraging in that region compared to the PCFG range. Both groups of gray whales spend the winter months on their breeding and calving grounds in Baja California, Mexico. 

Figure 1. The GEMM Lab field team following a Pacific Coast Feeding Group (PCFG) gray whale swimming in a kelp bed along the Oregon Coast during the summer foraging season. 

In January 2019 an Unusual Mortality Event (UME) was declared for gray whales due to the elevated numbers of stranded gray whales between Mexico and the Arctic regions of Alaska. Most of the stranded whales were emaciated, indicating that reduced nutrition and starvation may have been the causal factor of death. It is estimated that the population dropped from ~27,000 individuals in 2016 to ~21,000 in 2020 (Stewart & Weller, 2021).

During this UME period, between 2017-2019, the GEMM Lab was using drones to monitor the body condition of PCFG gray whales on their Oregon coastal feeding grounds (Fig. 1), while Christiansen and colleagues (2020) was using drones to monitor gray whales on their breeding grounds in San Ignacio Lagoon (SIL) in Baja California, Mexico. We teamed up with Christiansen and colleagues to compare the body condition of gray whales in these two different areas leading up to the UME. Comparing the body condition between these two populations could help inform which population was most effected by the UME.

The combined datasets consisted of four different drones used, thus different levels of photogrammetric uncertainty to consider. The GEMM Lab collected data using a DJI Phantom 3 Pro, DJI Phantom 4, and DJI Phantom 4 Pro, while Christiansen et al., (2020) used a DJI Inspire 1 Pro. By using the methodological approach described in my previous blog (here, also see Bierlich et al., 2021a for more details), we quantified photogrammetric uncertainty specific to each drone, allowing cross-comparison between these datasets. We also used Body Area Index (BAI), which is a standardized relative measure of body condition developed by the GEMM Lab (Burnett et al., 2018) that has low uncertainty with high precision, making it easier to detect smaller changes between individuals (see blog here, Bierlich et al., 2021b). 

While both PCFG and ENP gray whales visit San Ignacio Lagoon in the winter, we assume that the photogrammetry data collected in the lagoon is mostly of ENP whales based on their considerably higher population abundance. We also assume that gray whales incur low energetic cost during migration, as gray whale oxygen consumption rates and derived metabolic rates are much lower during migration than on foraging grounds (Sumich, 1983). 

Interestingly, we found that gray whale body condition on their wintering grounds in San Ignacio Lagoon deteriorated across the study years leading up to the UME (2017-2019), while the body condition of PCFG whales on their foraging grounds in Oregon concurrently increased. These contrasting trajectories in body condition between ENP and PCFG whales implies that dynamic oceanographic processes may be contributing to temporal variability of prey available in the Arctic/sub-Arctic and PCFG range. In other words, environmental conditions that control prey availability for gray whales are different in the two areas. For the ENP population, this declining nutritive gain may be associated with environmental changes in the Arctic/sub-Arctic region that impacted the predictability and availability of prey. For the PCFG population, the increase in body condition across years may reflect recovery of the NE Pacific Ocean from the marine heatwave event in 2014-2016 (referred to as “The Blob”) that resulted with a period of low prey availability. These findings also indicate that the ENP population was primarily impacted in the die-off from the UME. 

Surprisingly, the body condition of PCFG gray whales in Oregon was regularly and significantly lower than whales in San Ignacio Lagoon (Fig. 2). To further investigate this potential intrinsic difference in body condition between PCFG and ENP whales, we compared opportunistic photographs of gray whales feeding in the Northeastern Chukchi Sea (NCS) in the Arctic collected from airplane surveys. We found that the body condition of PCFG gray whales was significantly lower than whales in the NCS, further supporting our finding that PCFG whales overall have lower body condition than ENP whales that feed in the Arctic (Fig. 3). 

Figure 2. Boxplots showing the distribution of Body Area Index (BAI) values for gray whales imaged by drones in San Ignacio Lagoon (SIL), Mexico and Oregon, USA. The data is grouped by phenology group: End of summer feeding season (departure Oregon vs. arrival SIL) and End of wintering season (arrival Oregon vs. departure SIL). The group median (horizontal line), interquartile range (IQR, box), maximum and minimum 1.5*IQR (vertical lines), and outliers (dots) are depicted in the boxplots. The overlaid points represent the mean of the posterior predictive distribution for BAI of an individual and the bars represents the uncertainty (upper and lower bounds of the 95% HPD interval). Note how PCFG whales at then end of the feeding season (dark green) typically have lower body condition (as BAI) compared to ENP whales at the end of the feeding season when they arrive to SIL after migration (light brown).
Figure 3. Boxplots showing the distribution of Body Area Index (BAI) values of gray whales from opportunistic images collected from a plane in Northeaster Chukchi Sea (NCS) and from drones collected by the GEMM Lab in Oregon. The boxplots display the group median (horizontal line), interquartile range (IQR box), maximum and minimum 1.5*IQR (vertical lines), and outlies (dots). The overlaid points are the BAI values from each image. Note the significantly lower BAI of PCFG whales on Oregon feeding grounds compared to whales feeding in the Arctic region of the NCS.

This difference in body condition between PCFG and ENP gray whales raises some really interesting and prudent questions. Does the lower body condition of PCFG whales make them less resilient to changes in prey availability compared to ENP whales, and thus more vulnerable to climate change? If so, could this influence the reproductive capacity of PCFG whales? Or, are whales that recruit into the PCFG adapted to a smaller morphology, perhaps due to their specialized foraging tactics, which may be genetically inherited and enables them to survive with reduced energy stores?

These questions are on our minds here at the GEMM Lab as we prepare for our seventh consecutive field season using drones to collect data on PCFG gray whale body condition. As discussed in a previous blog by Dr. Alejandro Fernandez Ajo, we are combining our sightings history of individual whales, fecal hormone analyses, and photogrammetry-based body condition to better understand gray whales’ reproductive biology and help determine what the consequences are for these PCFG whales with lower body condition.

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Bierlich, K. C., Hewitt, J., Bird, C. N., Schick, R. S., Friedlaender, A., Torres, L. G., … & Johnston, D. W. (2021). Comparing Uncertainty Associated With 1-, 2-, and 3D Aerial Photogrammetry-Based Body Condition Measurements of Baleen Whales. Frontiers in Marine Science, 1729.

Bierlich, K. C., Schick, R. S., Hewitt, J., Dale, J., Goldbogen, J. A., Friedlaender, A.S., et al. (2021b). Bayesian Approach for Predicting Photogrammetric Uncertainty in Morphometric Measurements Derived From Drones. Mar. Ecol. Prog. Ser. 673, 193–210. doi: 10.3354/meps13814

Burnett, J. D., Lemos, L., Barlow, D., Wing, M. G., Chandler, T., & Torres, L. G. (2018). Estimating morphometric attributes of baleen whales with photogrammetry from small UASs: A case study with blue and gray whales. Marine Mammal Science35(1), 108–139.

Christiansen, F., Rodrı́guez-González, F., Martı́nez-Aguilar, S., Urbán, J., Swartz, S., Warick, H., et al. (2021). Poor Body Condition Associated With an Unusual Mortality Event in Gray Whales. Mar. Ecol. Prog. Ser. 658, 237–252. doi:10.3354/meps13585

Hildebrand, L., Bernard, K. S., and Torres, L. G. (2021). Do Gray Whales Count Calories? Comparing Energetic Values of Gray Whale Prey Across Two Different Feeding Grounds in the Eastern North Pacific. Front. Mar. Sci. 8. doi: 10.3389/fmars.2021.683634

Stewart, J. D., and Weller, D. (2021). Abundance of Eastern North Pacific Gray Whales 2019/2020 (San Diego, CA: NOAA/NMFS)

Sumich, J. L. (1983). Swimming Velocities, Breathing Patterns, and Estimated Costs of Locomotion in Migrating Gray Whales, Eschrichtius Robustus. Can. J. Zoology. 61, 647–652. doi: 10.1139/z83-086

Torres, L.G., Bird, C., Rodrigues-Gonzáles, F., Christiansen F., Bejder, L., Lemos, L., Urbán Ramírez, J., Swartz, S., Willoughby, A., Hewitt., J., Bierlich, K.C. (2022). Range-wide comparison of gray whale body condition reveals contrasting sub-population health characteristics and vulnerability to environmental change. Frontiers in Marine Science. 9:867258.

Dive into Oregon’s underwater forests

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

When I was younger, I aspired to be a marine mammal biologist. I thought it was purely about knowing as much about marine mammal species as possible. However, over time and with experience in this field, I have realized that in order to understand a species, you need to have a holistic understanding of its prey, habitat, and environment. When I first applied to be advised by Leigh in the GEMM Lab, I had no idea how much of my time I would spend looking at tiny zooplankton under a microscope, thinking about the different benefits of different habitat types, or reading about oceanographic processes. But these things have been incredibly vital to my research to date and as a result, I now refer to myself as a marine ecologist. This holistic understanding that I am gaining will only grow throughout my PhD as I am broadly looking at the habitat use, site fidelity, and population dynamics of the Pacific Coast Feeding Group (PCFG) of gray whales for my thesis research. 

The PCFG display many foraging tactics and occupy several habitat types along the Oregon coast while they spend their summer feeding seasons here (Torres et al. 2018). Here, I will focus on one of these habitats: kelp. When you hear the word kelp, you probably conjure an image of long, thick stalks that reach from the ocean floor to the surface, with billowing fronds waving around (Figure 1a). However, this type is only one of three basic morphologies (Filbee-Dexter & Scheibling 2014) and it is called canopy kelp, which often forms extensive forests. The other two morphologies are stipitate and prostrate kelps. The former forms midwater stands (Figure 1b) while the latter forms low-lying kelp beds (Figure 1c). All three of these morphologies exist on the Oregon coast and create a mosaic of understory and canopy kelp patches that dot our coastline.

Figure 1. Examples of the three different kelp morphologies. a: bull kelp (Nereocystis luetkeana) is a type of canopy kelp and the dominant kelp on the Oregon coast (Source: Oregon Coast Aquarium); b: sea palm (Postelsia palmaeformis) is a type of stipitate kelp that forms mid-water stands (Source: Oregon Conservation Strategy); c: sea cabbage (Saccharina sessilis) is a type of prostrate kelp that is stipeless and forms low-lying kelp beds (Source: Central Coast Biodiversity).

One of the most magnificent things about kelp is that it is not just a species itself, but it provides critical habitat, refuge, and food resources to a myriad of other species due to its high rates of primary production (Dayton 1985). Kelp is often referred to as a foundation species due to all of these critical services it provides. In Oregon, many species of rockfish, which are important commercial and recreational fisheries, use kelp as habitat throughout their life cycle, including as nursery grounds. Lingcod, another widely fished species, forages amongst kelp. A large number of macroinvertebrates can be found in Oregon kelp forests, including anemones, limpets, snails, sea urchins, sea stars, and abalone, to name a fraction of them. 

Kelps grow best in cold, nutrient-rich waters (Tegner et al. 1996) and their growth and distribution patterns are highly naturally variable on both temporal and spatial scales (Krumhansl et al. 2016). However, warm water, low nutrient or light conditions, intensive grazing by herbivores, and severe storm activity can lead to the erosion and defoliation of kelp beds (Krumhansl et al. 2016). While these events can occur naturally in cyclical patterns, the frequency of several of these events has increased in recent years, as a result of climate change and anthropogenic impacts. For example, Dawn’s blog discussed increasing marine heatwaves that represent an influx of warm water for a prolonged period of time. In fact, kelps can be useful sentinels of change as they tend to be highly responsive to changes in environmental conditions (e.g., Rogers-Bennet & Catton 2019) and their nearshore, coastal location directly exposes them to human activities, such as pollution, harvesting, and fishing (Bennett et al. 2016).

Due to its foundational role, changes or impacts to kelp can reverberate throughout the ecosystem and negatively affect many other species. As mentioned previously, kelp is naturally highly variable, and like many other ecological processes, undergoes boom and bust cycles. For over four decades, dense, productive kelp forests have been shown to transition to sea urchin barrens, and back again, in natural cycles (Sala et al. 1998; Pinnegar et al. 2000; Steneck et al. 2002; Figure 2). These transitions are called phase shifts. In a healthy, balanced kelp forest, sea urchins typically passively feed on detrital plant matter, such as broken off pieces of kelp fronds that fall to the seafloor. A phase shift occurs when the grazing intensity of sea urchins increases, resulting in them actively feeding on kelp stalks and fronds to a point where the kelp in an area can become greatly reduced, creating an urchin barren. Sea urchin grazing intensity can change for a number of reasons, including reduction in sea urchin predators (e.g., sea otters, sunflower sea stars) or poor kelp recruitment events (e.g., due to warm water temperature). Regardless of the reason, the phases tend to transition back and forth over time. However, there is concern that sea urchin barrens may become an alternative stable state of the subtidal ecosystem from which kelp in an area cannot recover (Filbee-Dexter & Scheibling 2014). 

Figure 2. Screenshots from GoPro videos from 2016 (left) and 2018 (right) at the same kayak sampling station in Port Orford showing the difference between a dense kelp forest and what appears to be an urchin barren. (Source: GEMM Lab).

For example, in 2014, bull kelp canopy cover in northern California was reduced by >90% and has not shown signs of recovery since (Rogers-Bennet & Catton 2019; Figure 3). This massive decline was attributed to two major events: 1) the onset of sea star wasting disease (SSWD) in 2013 and 2) the “warm blob” of 2014-2016. SSWD affected over 20 sea star species along the coast from Mexico to Alaska, with the predatory sunflower sea star, which consumes purple sea urchins, most affected, including population declines of 80-100% along the coast (Harvell et al. 2019). Following this SSWD outbreak, the “warm blob”, which was an extreme marine heatwave in the Pacific Ocean, caused ocean temperatures to spike. These two events allowed purple sea urchin populations to grow unchecked by their predators, and created nutrient-poor and warm water conditions, which limited kelp growth and productivity. Intense grazing on bull kelp by growing urchin populations resulted in the >90% reduction in bull kelp canopy cover and has left behind widespread urchin barrens instead (Rogers-Bennet & Catton 2019). Consequently, there have been ecological and economic impacts on the ecosystem and communities in northern California. Without bull kelp, red abalone and red sea urchin populations starved, leading to a subsequent loss of the recreational red abalone (estimated value of $44 million/year) and commercial red urchin fisheries in northern California (Rogers-Bennet & Catton 2019).

Figure 3. Surface kelp canopy area pre- and post-impact from sites in Sonoma and Mendocino counties, northern California from aerial surveys (2008, 2014-2016). Figure and figure caption taken from Rogers-Bennett & Catton (2019).

As I mentioned earlier, while phase shifts between kelp forests and urchin barrens are common cycles, the intensity of the events described above in northern California are an example of sea urchin barrens potentially becoming a stable state of the subtidal ecosystem (Filbee-Dexter & Scheibling 2014). Given that marine heatwaves are only expected to increase in intensity and frequency in the future (Frölicher et al. 2018), the events documented in northern California may not be an isolated incidence. 

Considering that parts of the Oregon coast, particularly the southern portion, are very similar to northern California biogeographically, and that it was not exempt from the “warm blob”, similar changes in kelp forests may be occurring along our coast. There are many individuals and groups that are actively working on this issue to examine potential impacts to kelp and the species that depend on the services it provides. For more information, check out the Oregon Kelp Alliance

Figure 4. A gray whale surfaces in a large kelp bed during a foraging bout along the Oregon coast. (Source: GEMM Lab).

So, what does all of this information have to do with gray whales? Given their affinity for kelp habitats (Figure 4) and their zooplankton prey that aggregates there, changes to kelp ecosystems may affect gray whale health and ecology. This aspect of the complex kelp trophic web has not been examined to date; thus one of my PhD chapters focuses on the response of gray whales to changing kelp ecosystems along the southern Oregon coast. To do this, I am examining 6 years of data collected during the TOPAZ/JASPER project in Port Orford, to look at the relationships between kelp health, sea urchin density, zooplankton abundance, and gray whale foraging effort over space and time. Documenting impacts of changing kelp forests on gray whales is important to assist management efforts as healthy and abundant kelp seems critical in providing ample food opportunities for these iconic Pacific Northwest marine predators.

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Bennett S, et al. The ‘Great Southern Reef’: Social, ecological and economic value of Australia’s neglected kelp forests. Marine and Freshwater Research 67:47-56.

Dayton PK (1985) Ecology of kelp communities. Annual Review of Ecology and Systematics 16:215-245.

Filbee-Dexter K, Scheibling RE (2014) Sea uechin barrens as alternative stable states of collapsed kelp ecosystems. Marine Ecology Progress Series 495:1-25.

Frölicher TL, Fischer EM, Gruber N (2018) Marine heatwaves under global warming. Nature 560:360-364.

Harvell CD, et al. (2019) Disease epidemic and a marine heat wave are associated with the continental-scale collapse of a pivotal predator (Pycnopodia helianthoides). Science Advances 5(1) doi:10.1126/sciadv.aau7042.

Krumhansl KA, et al. (2016) Global patterns of kelp forest change over the past half-century. Proceedings of the National Academy of Sciences of the United States of America 113(48):13785-13790.

Pinnegar JK, et al. (2000) Trophic cascades in benthic marine ecosystems: lessons for fisheries and protected-area management. Environmental Conservation 27:179-200.

Rogers-Bennett L, Catton CA (2019) Marine heat wave and multiple stressors tip bull kelp forest to sea urchin barrens. Scientific Reports 9:15050.

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A pregnancy test for whales?! Why and how?

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

I often receive two reactions when asked what I am currently working on; one is “Wow! That is a very cool job, it must be amazing to work with such incredible animals!”, the other is “How do you do that and why is that important?”. So, today I decided to blog about some of the reasons why it is important to develop a pregnancy test for gray whales and how we are doing this.

In a previous blogpost, I described the many ways in which whales play critical roles in sustaining marine ecosystem. Briefly, whales can enhance marine productivity by vertically and horizontally mixing of ocean waters, promoting primary production, and mitigating climate change by sequestering carbon with their large biomass and long life-span (1-3). Even after they die, their carcasses can contribute to biodiversity creating new habitat on the seafloor (4). But, over several decades, the whaling industry drastically removed whales around the globe, with some species and populations depleted to near extinction (5). Consequently, these depleted whale populations now play a diminished role in ocean ecosystem processes and their recovery is currently challenged by an increasing number of modern anthropogenic impacts. Hence, working towards whale conservation is essential for keeping a healthy marine ecosystem.

Working and designing effective strategies for conservation biology often involves gaining knowledge regarding the reproductive parameters of individual animals in wild populations. This information is critical for understanding population trends and the underlaying mechanisms that affect animal welfare and their potential for recovery. However, getting such information from free-living whales can be challenging (see Hunt et al. 2013). While we know that whales typically have long life-spans, lengthy generation times, extended parental care, and high survival rates, detailed knowledge on the life history and general reproductive biology of free-ranging whales is limited for the majority of the whale populations. In fact, much of what we do know about whale reproduction is derived from whaling records. Only recently, conservation physiology approaches (see our previous post here) have contributed alternative and non-invasive methods for monitoring key physiological processes that can help monitor a whale’s reproductive biology and determine reproductive parameters such as sexual maturity and pregnancy (6-9).

In this clip you can see an example of a fecal sample collection from a gray whale off the Oregon coast. We can look at hormones in the fecal samples which are useful indicators for endocrine assessments of free-swimming whales. Fecal sample and footage filmed under NOAA/NMFS permit #16111.

Gray whales (Eschrichtius robustus) in the Eastern North Pacific (ENP) typically undertake annual migrations between their lower latitude breeding grounds in the coastal waters of the Baja California Peninsula, Mexico, and the foraging grounds located on the Bering and Chukchi Seas (10). However, among the ENP whales a distinct subgroup of about 230 whales shorten their migration to feed in the coastal waters of Northern California, Oregon, and southeastern Alaska (11). This group of whales is known as the gray whale Pacific Coast Feeding Group (PCFG).

Since 2016, the GEMM Lab has monitored individual gray whales within the PCFG off the Oregon coast (check the GRANITE project). Gray whales have a distinct mottled skin; and each individual whale presents a unique pigmentation pattern that allows for the individual identification of whales. We can identify who is who among the whales who visit the Oregon coast. In this way, we can keep a detailed record of re-sightings of known individuals (visit our new web site to know more about the lives of individual whales that visit the Oregon coast).  We have high individual re-sighting rates, so this unique opportunity helps us keep a long-term data series for individual whales to monitor their health, body condition, and reproductive status over time, and thus further develop and advance our non-invasive study methods.

We are combining behavioral and feeding ecology with drone photogrammetry and endocrinology of the same individual whales to help us understand the relationships between natural and anthropogenic drivers with biological parameters. In this way, following individual whales, we are developing sensitive biomarkers to monitor and infer about the population health, population trends, and identify stressors that impact their recovery and welfare. In particular, we are now working to develop a noninvasive approach to detect pregnancy in gray whales based on fecal hormone analyses.

In this picture you can see “Rose”, a gray whale calf, on top of her mother “Scarlett”. Scarlett is one of the most recognizable whales from the PCFG, due to a large scar on the right side of her back (not visible in this picture). She has been observed along the Pacific NW coast since 1996, so she is at least 26 years old today. We know 3 of her calves. Following individual whales like Scarlett is helping us to better understand the gray whale reproductive biology. Photo by Alejandro Fernandez Ajo taken under NOAA/NMFS permit #21678.

In marine mammals, the progesterone hormone is secreted in the ovaries during the estrous cycle and gestation, and is the predominant hormone responsible for sustaining pregnancy (12). As the hormones are cleared from the blood into the gut, they are metabolized and eventually excreted in feces; fecal samples represent a cumulative and integrated concentration of hormone metabolites (13-14), which are useful indicators for endocrine assessments of free-swimming whales. Several studies show that changes in hormone concentration correlate in meaningful ways with exposure to stressors (15-16) and changes in reproductive status (17-19). We are using our long data series of fecal hormones and individual life histories to advance our understanding on the gray whales’ reproductive biology. We are close to developing a technique that will allow us to detect pregnancy in whales based in fecal hormones analyses and photogrammetry. Stay tuned for results from this pregnancy test!

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1- Pershing AJ, Christensen LB, Record NR, Sherwood GD, Stetson PB (2010) The impact of whaling on the ocean carbon cycle: Why bigger was better. PLoS ONE 5(8): e12444.

2- Roman J and McCarthy JJ. 2010. The whale pump: marine mammals enhance primary productivity in a coastal basin. PLoS ONE. 5(10): e13255.

3- Morissette L, Kaschner K, and Gerber LR. 2010. “Whales eat fish”? Demystifying the myth in the Caribbean marine ecosystem. Fish Fish 11: 388–404.

4- Smith CR, Roman J, Nation JB. A metapopulation model for whale-fall specialists: The largest whales are essential to prevent species extinctions. J. Mar. Res. 77, 283–302 (2019).

5- Branch TA, Williams TM. Legacy of industrial whaling. Whales. Whal. Ocean Ecosyst. 2006, 262–278 (2006).

6- Kellar NM, Keliher J, Trego ML, Catelani KN, Hanns C, George JC, et al. Variation of bowhead whale progesterone concentrations across demographic groups and sample matrices. Endanger Species Res 2013; 22:61–72.

7- Pallin L, Robbins J, Kellar N, Berube M, Friedlaender A. Validation of a blubber-based endocrine pregnancy test for humpback whales. Conserv Physiol 2018;6:1 11.

8-Hunt KE, Robbins J, Buck CL, Bérubé M, Rolland RM (2019) Evaluation of fecal hormones for noninvasive research on reproduction and stress in humpback whales (Megaptera novaeangliae). Gen Comp Endocrinol 280: 24–34.

9-Melica, V., Atkinson, S., Calambokidis, J., Lang, A., Scordino, J., & Mueter, F. (2021). Application of endocrine biomarkers to update information on reproductive physiology in gray whale (Eschrichtius robustus). Plos one, 16(8), e0255368.

10-Swartz SL. Gray Whale. In: Wursig B, Thewissen JGM, Kovacs KM, editors. Encyclopedia of Marine Mammals (Third Edition). Elsevier;2018,p. 422–8.–0.

11-Calambokidis J, Darling JD, Deecke V, Gearin P, Gosho M, Megill W, et al. Abundance, range and movements of a feeding aggregation of gray whales (Eschrichtius robustus) from California to south-eastern Alaska in 1998. J Cetacean Res Manag 2002;4:267–76.

12- Bronson, F. H. (1989). Mammalian reproductive biology. University of Chicago Press.

13-Wasser SK, Hunt KE, Brown JL, Cooper K, Crockett CM, Bechert U, Millspaugh JJ, Larson S, Monfort SL (2000) A generalized fecal glucocorticoid assay for use in a diverse array of nondomestic mammalian and avian species. Gen Comp Endocrinol120:260–275.

14- Hunt, K.E., Rolland, R.M., Kraus, S.D., Wasser, S.K., 2006. Analysis of fecal glucocorticoids in the North Atlantic right whale (Eubalaena glacialis). Gen. Comp. Endocrinol. 148, 260–272.

15- Lemos, L.S., Olsen, A., Smith, A., Burnett, J.D., Chandler, T.E., Larson, S., Hunt, K.E., Torres, L.G., 2021. Stressed and slim or relaxed and chubby? A simultaneous assessment of gray whale body condition and hormone variability. Mar. Mammal Sci. 1–11.

16- Rolland, R., McLellan, W., Moore, M., Harms, C., Burgess, E., Hunt, K., 2017. Fecal glucocorticoids and anthropogenic injury and mortality in North Atlantic right whales Eubalaena glacialis. Endanger. Species Res. 34, 417–429.

17-Rolland, R.M., Hunt, K.E., Kraus, S.D., Wasser, S.K., 2005. Assessing reproductive status of right whales (Eubalaena glacialis) using fecal hormone metabolites. Gen. Comp. Endocrinol. 142, 308–317.

18- Valenzuela Molina M, Atkinson S, Mashburn K, Gendron D, Brownell RL. Fecal steroid hormones reveal reproductive state in female blue whales sampled in the Gulf of California, Mexico. Gen Comp Endocrinol 2018;261:127–35. PMID:29476760.

19- Hunt, K. E., Robbins, J., Buck, C. L., Bérubé, M., & Rolland, R. M. (2019). Evaluation of fecal hormones for noninvasive research on reproduction and stress in humpback whales (Megaptera novaeangliae). General and Comparative Endocrinology, 280, 24-34.