Leigh Torres – Geospatial Ecology of Marine Megafauna Laboratory

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

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

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

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

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

A blue whale comes up for air in the South Taranaki Bight. Photo by Leigh Torres.

Project goals

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

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

Kicking off fieldwork

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

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

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

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

Research team

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

Principal Investigators:

Leigh Torres (Geospatial Ecology of Marine Megafauna Lab, Marine Mammal Institute, OSU)

Research partners and key collaborators:

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References:

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

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

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

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

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

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

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

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

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

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

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

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

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

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

*“Lunita”, an example of successful internal recruitment*

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

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

References

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

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

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

Learning by listening

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

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

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

The South Taranaki Bight: an ideal study system

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

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

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

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

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

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

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

Marine heatwaves impact multiple life history processes

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

More than a decade of research on New Zealand blue whales

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

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

References:

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

References

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

References

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

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

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

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

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

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

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

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

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

Spitz J, Trites AW, Becquet V, Brind’Amour A, Cherel Y, Galois R, Ridoux V (2012) Cost of living dictates what whales, dolphins and porpoises eat: the importance of prey quality on predator foraging strategies. PLOS ONE 7:e50096

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

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

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

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

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

Introduction

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.*

Expectations

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.*

Conclusion

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.

References

Benoit-Bird, K. J., & McManus, M. A. (2012). Bottom-up regulation of a pelagic community through spatial aggregations. Biology Letters, 8(5), 813–816. https://doi.org/10.1098/rsbl.2012.0232

Burnett, J. D., & Wing, M. G. (2018). A low-cost near-infrared digital camera for fire detection and monitoring. International Journal of Remote Sensing, 39(3), 741–753. https://doi.org/10.1080/01431161.2017.1385109

Sinclair, A. R. E., & Krebs, C. J. (2002). Complex numerical responses to top–down and bottom–up processes in vertebrate populations. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 357(1425), 1221–1231.https://doi.org/10.1098/rstb.2002.1123.

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 8^th 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 24^th. 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 24^th 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.

*Porcelain crab larvae. Source: GEMM Lab.*

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

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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Reference

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.

Acknowledgments

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.

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.

References

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.

Sala E, Boudouresque CF, Harmelin-Vivien M (1998) Fishing, trophic cascades and the structure of algal assemblages; evaluation of an old but untested paradigm. Oikos 82:425-439.

Steneck RS, et al. (2002) Kelp forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation 29:436-459.

Tegner MJ, Dayton PK, Edwards PB, Riser KL (1996) Is there evidence for the long-term climatic change in southern California kelp forests? California Cooperative Oceanic Fisheries Investigations Report 37:111-126.

Torres LG, Nieukirk SL, Lemos L, Chandler TE (2018) Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science doi:10.3389/fmars.2019.00319.

Wavelet analysis to describe biological cycles and signals of non-stationarity

By Allison Dawn, GEMM Lab Master’s student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

During my second term of graduate school, I have been preparing to write my research proposal. The last two months have been an inspiring process of deep literature dives and brainstorming sessions with my mentors. As I discussed in my last blog, I am interested in questions related to pattern and scale (fine vs. mesoscale) in the context of the Pacific Coast Feeding Group (PCFG) of gray whales, their zooplankton prey, and local environmental variables.

My work currently involves exploring which scales of pattern are important in these trophic relationships and whether the dominant scale of a pattern changes over time or space. I have researched which analysis tools would be most appropriate to analyze ecological time series data, like the impressive long-term dataset the GEMM lab has collected in Port Orford as part of the TOPAZ project, where we have monitored the abundance of whales and zooplankton, as well as environmental variables since 2016.

A useful analytical tool that I have come across in my recent coursework and literature review is called wavelet analysis. Importantly, wavelet analysis can handle non-stationarity and edge detection in time series data. Non-stationarity is when a dataset’s mean and/or variance can change over time or space, and edge detection is the identification of the change location (in time or space). For example, it is not just the cycles or “ups and downs” of zooplankton abundance I am interested in, but when in time or where in space these cycles of “ups and downs” might change in relation to what their previous values, or distances between values, were. Simply stated, non-stationarity is when what once was normal is no longer normal. Wavelet analysis has been applied across a broad range of fields, such as environmental engineering (Salas et al. 2020), climate science (Slater et al. 2021), and bio-acoustics (Buchan et al. 2021). It can be applied to any time series dataset that might violate the traditional statistical assumption of stationarity.

In a recent review of climate science methodology, Slater et al. (2021) outlined the possible behavior of time series data. Using theoretical plots, the authors show that data can a) have the same mean and variance over time, or b) have non-stationarity that can be broken into three major groups – trend, step change, or shifts in variance. Figure 1 further demonstrates the difference between stationary vs. non-stationary data in relation to a given variable of interest over time.

Figure 1. Plots showing the possible magnitude of a given variable across a time series: a) Stationary behavior, b) Non-stationary trend, step-change, and a shift in variance. [Taken from Slater et. al (2021)].

Traditional correlation statistics assumes stationarity, but it has been shown that ecological time series are often non-stationary at certain scales (Cazelles & Hales, 2006). In fact, ecological data rarely meets the requirements of a controlled experiment that traditional statistics require. This non-stationarity of ecological data means that while widely-used methods like generalized linear models and analyses of variances (ANOVAs) can be helpful to assess correlation, they are not always sufficient on their own to describe the complex natural phenomena ecologists seek to explain. Non-stationarity occurs frequently in ecological time series, so it is appropriate to consider analysis tools that will allow us to detect edges to further investigate the cause.

Wavelet analysis can also be conducted across a time series of multiple response variables to assess if these variables share high common power (correlation). When data is combined in this way it is called a cross-wavelet analysis. An interesting paper used cross-wavelet analysis to assess the seasonal response of zooplankton life history in relation to climate warming (Winder et. al 2009). Results from their cross-wavelet analysis showed that warming temperatures over the past two decades increased the voltinism (number of broods per year) of copepods. The authors show that where once annual recruitment followed a fairly stationary pattern, climate warming has contributed to a much more stochastic pattern of zooplankton abundance. From these results, the authors contribute to the hypothesis that climate change has had a temporal impact on zooplankton population dynamics, and recruitment has increasingly drifted out of phase from the original annual cycles.

Figure 2. Cross-wavelet spectrum for immature and adult Leptodiaptomus ashlandi for 1965 through either 2000 or 2005. Plots show a) immatures and temperature, b) adults and temperature, c) immatures and phytoplankton, and d) adults and phytoplankton. Arrows indicate phase between combined time series. 0 degrees is in-phase and 180 degrees is anti-phase. Black contour lines show “cone of influence” or the 95% significance level, every value within the cone is considered significant. Left axis shows the temporal period, and the color legend shows wavelet frequency power, with low frequencies in blue and high frequencies in red. Plots show strong covariation of high common power at the 12-month period until the 1980s. This pattern is especially evident in plot c) and d). [Taken from (Winder et. al 2009)].

While wavelet and cross-wavelet analyses should not be the only tool used to explore data, due to its limitations with significance testing, it is still worth implementing to gain a better understanding of how time series variables relate to each other over multiple spatial and/or temporal scales. It is often helpful to combine multiple methods of analysis to get a larger sense of patterns in the data, especially in spatio-temporal research.

When conducting research within the context of climate change, where the concentration of CO₂ in ppm in the atmosphere is a non-stationary time series itself (Figure 3), it is important to consider how our datasets might be impacted by climate change and wavelet analysis can help identify the scales of change.

Figure 3. Plot showing the historic fluctuations of CO^2 and the recent deviation from normal levels. Source: https://globalclimate.ucr.edu/resources.html

When considering our ecological time series of data in Port Orford, we want to evaluate how changing ocean conditions may be related to data trends. For example, has the annual mean or variance of zooplankton abundance changed over time, and where has that change occurred in time or space? These changes might have occurred at different scales and might be invisible at other scales. I am eager to see if wavelet analysis can detect these sorts of changes in the abundance of zooplankton across our time series of data, particularly during the seasons of intense heat waves or upwelling.

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References

Buchan, S. J., Pérez-Santos, I., Narváez, D., Castro, L., Stafford, K. M., Baumgartner, M. F., … & Neira, S. (2021). Intraseasonal variation in southeast Pacific blue whale acoustic presence, zooplankton backscatter, and oceanographic variables on a feeding ground in Northern Chilean Patagonia. Progress in Oceanography, 199, 102709.

Cazelles, B., & Hales, S. (2006). Infectious diseases, climate influences, and nonstationarity. PLoS Medicine, 3(8), e328.

Salas, J. D., Anderson, M. L., Papalexiou, S. M., & Frances, F. (2020). PMP and climate variability and change: a review. Journal of Hydrologic Engineering, 25(12), 03120002.

Slater, L. J., Anderson, B., Buechel, M., Dadson, S., Han, S., Harrigan, S., … & Wilby, R. L. (2021). Nonstationary weather and water extremes: a review of methods for their detection, attribution, and management. Hydrology and Earth System Sciences, 25(7), 3897-3935.

Winder, M., Schindler, D. E., Essington, T. E., & Litt, A. H. (2009). Disrupted seasonal clockwork in the population dynamics of a freshwater copepod by climate warming. Limnology and Oceanography, 54(6part2), 2493-2505.

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