As waters warm, what are “anomalous conditions” in the face of climate change?

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

Recently, I had the opportunity to attend the Effects of Climate Change on the World’s Ocean (ECCWO) conference. This meeting brought together experts from around the world for one week in Bergen, Norway, to gather and share the latest information on how oceans are changing, what is at risk, responses that are underway, and strategies for increasing climate resilience, mitigation, and adaptation. I presented our recent findings from the EMERALD project, which examines gray whale and harbor porpoise distribution in the Northern California Current over the past three decades. Beyond sharing my postdoctoral research widely for the first time and receiving valuable feedback, the ECCWO conference was an incredibly fruitful learning experience. Marine mammals can be notoriously difficult to study, and often the latest methodological approaches or conceptual frameworks take some time to make their way into the marine mammal field. At ECCWO, I was part of discussions at the ground floor of how the scientific community can characterize the impacts of climate change on the ecosystems, species, and communities we study.

One particular theme became increasingly apparent to me throughout the conference: as the oceans warm, what are “anomalous conditions”? There was an interesting dichotomy between presentations focusing on “extreme events,” “no-analog conditions,” or “non-stationary responses,” compared with discussions about the overall trend of increasing temperatures due to climate change. Essentially, the question that kept arising was, what is our frame of reference? When measuring change, how do we define the baseline?

Marine heatwaves have emerged as an increasingly prevalent phenomenon in recent years (see previous GEMM Lab blogs about marine heatwaves here and here). The currently accepted and typically applied definition of a marine heatwave is when water temperatures exceed a seasonal threshold (greater than the 90th percentile) for a given length of time (five consecutive days or longer) (Hobday et al. 2016). These marine heatwaves can have substantial ecosystem-wide impacts including changes in water column structure, primary production, species composition, distribution, and health, and fisheries management such as closures and quota changes (Cavole et al. 2016, Oliver et al. 2018). Through some of our own previous research, we documented that blue whales in Aotearoa New Zealand shifted their distribution (Barlow et al. 2020) and reduced their reproductive effort (Barlow et al. 2023) in response to marine heatwaves. Concerningly, recent projections anticipate an increase in the frequency, intensity, and duration of marine heatwaves under global climate change (Frölicher et al. 2018, Oliver et al. 2018).

However, as the oceans continue to warm, what baseline do we use to define anomalous events like marine heatwaves? Members of the US National Oceanic and Atmospheric Administration (NOAA) Marine Ecosystem Task Force recently put forward a comment article in Nature, proposing revised definitions for marine heatwaves under climate change, so that coastal communities have the clear information they need to adapt (Amaya et al. 2023). The authors posit that while a “fixed baseline” approach, which compares current conditions to an established period in the past and has been commonly used to-date (Hobday et al. 2016), may be useful in scenarios where a species’ physiological limit is concerned (e.g., coral bleaching), this definition does not incorporate the combined effect of overall warming due to climate change. A “shifting baseline” approach to defining marine heatwaves, in contrast, uses a moving window definition for what is considered “normal” conditions. Therefore, this shifting baseline approach would account for long-term warming, while also calculating anomalous conditions relative to the current state of the system.

An overview of two different definitions for marine heatwaves, relative to either fixed or shifting baselines. Reproduced from Amaya et al. 2023.

Why bother with these seemingly nuanced definitions and differences in terminology, such as fixed versus shifting baselines for defining marine heatwave events? The impacts of these events can be extreme, and potentially bear substantial consequences to ecosystems, species, and coastal communities that rely on marine resources. With the fixed baseline definition, we may be headed toward perpetual heatwave conditions (i.e., it’s almost always hotter than it used to be), at which point disentangling the overall warming trends from these short-term extremes becomes nearly impossible. What the shifting baseline definition means in practice, however, is that in the future temperatures would need to be substantially higher than the historical average in order to qualify as a marine heatwave, which could obscure public perception from the concerning reality of warming oceans. Yet, the authors of the Nature comment article claim, “If everything is extremely warm all of the time, then the term ‘extreme’ loses its meaning. The public might become desensitized to the real threat of marine heatwaves, potentially leading to inaction or a lack of preparedness.” Therefore, clear messaging surrounding both long-term warming and short-term anomalous conditions are critically important for adaptation and resource allocation in the face of rapid environmental change.

While the findings presented and discussed at an international climate change conference could be considered quite disheartening, I left the ECCWO conference feeling re-invigorated with hope. Crown Prince Haakon of Norway gave the opening plenary and articulated that “We need wise and concerned scientists in our search for truth”. Later in the week, I was a co-convenor of a session that gathered early-career ocean professionals, where we discussed themes such as how we deal with uncertainty in our own climate change-related ocean research, and importantly, how do we communicate our findings effectively. Throughout the meeting, I had formal and informal discussions about methods and analytical techniques, and also about what connects each of us to the work that we do. Interacting with driven and dedicated researchers across a broad range of disciplines and career stages gave me some renewed hope for a future of ocean science and marine conservation that is constructive, collaborative, and impactful.

Enjoying the ~anomalously~ sunny April weather in Bergen, Norway, during the ECCWO conference.

Now, as I am diving back in to understanding the impacts of environmental conditions on harbor porpoise and gray whale habitat use patterns through the EMERALD project, I am keeping these themes and takeaways from the ECCWO conference in mind. The EMERALD project draws on a dataset that is about as old as I am, which gives me some tangible perspective on how things have things changed in the Northern California Current during my lifetime. We are grappling with what “anomalous” conditions are in this dynamic upwelling system on our doorstep, whether these anomalies are even always bad, and how conditions continue to change in terms of cyclical oscillations, long-term trends, and short-term events. Stay tuned for what we’ll find, as we continue to disentangle these intertwined patterns of change.

Did you enjoy this blog? Want to learn more about marine life, research, and conservation? Subscribe to our blog and get a weekly alert when we make a new post! Just add your name into the subscribe box below!



Amaya DJ, Jacox MG, Fewings MR, Saba VS, Stuecker MF, Rykaczewski RR, Ross AC, Stock CA, Capotondi A, Petrik CM, Bograd SJ, Alexander MA, Cheng W, Hermann AJ, Kearney KA, Powell BS (2023) Marine heatwaves need clear definitions so coastal communities can adapt. Nature 616:29–32.

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, 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.

Cavole LM, Demko AM, Diner RE, Giddings A, Koester I, Pagniello CMLS, Paulsen ML, Ramirez-Valdez A, Schwenck SM, Yen NK, Zill ME, Franks PJS (2016) Biological impacts of the 2013–2015 warm-water anomaly in the northeast Pacific: Winners, losers, and the future. Oceanography 29:273–285.

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

Hobday AJ, Alexander L V., Perkins SE, Smale DA, Straub SC, Oliver ECJ, Benthuysen JA, Burrows MT, Donat MG, Feng M, Holbrook NJ, Moore PJ, Scannell HA, Sen Gupta A, Wernberg T (2016) A hierarchical approach to defining marine heatwaves. Prog Oceanogr.

Oliver ECJ, Donat MG, Burrows MT, Moore PJ, Smale DA, Alexander L V., Benthuysen JA, Feng M, Sen Gupta A, Hobday AJ, Holbrook NJ, Perkins-Kirkpatrick SE, Scannell HA, Straub SC, Wernberg T (2018) Longer and more frequent marine heatwaves over the past century. Nat Commun 9:1–12.

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.

Did you enjoy this blog? Want to learn more about marine life, research, and conservation? Subscribe to our blog and get a weekly alert when we make a new post! Just add your name into the subscribe box below!



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.

Marine megafauna as ecosystem sentinels: What animals can tell us about changing oceans

By Dawn Barlow1 and Will Kennerley2

1PhD Candidate, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

2MS Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Seabird Oceanography Lab

The marine environment is dynamic, and mobile animals must respond to the patchy and ephemeral availability of resource in order to make a living (Hyrenbach et al. 2000). Climate change is making ocean ecosystems increasingly unstable, yet these novel conditions can be difficult to document given the vast depth and remoteness of most ocean locations. Marine megafauna species such as marine mammals and seabirds integrate ecological processes that are often difficult to observe directly, by shifting patterns in their distribution, behavior, physiology, and life history in response to changes in their environment (Croll et al. 1998, Hazen et al. 2019). These mobile marine animals now face additional challenges as rising temperatures due to global climate change impact marine ecosystems worldwide (Hazen et al. 2013, Sydeman et al. 2015, Silber et al. 2017, Becker et al. 2019). Given their mobility, visibility, and integration of ocean processes across spatial and temporal scales, these marine predator species have earned the reputation as effective ecosystem sentinels. As sentinels, they have the capacity to shed light on ecosystem function, identify risks to human health, and even predict future changes (Hazen et al. 2019). So, let’s explore a few examples of how studying marine megafauna has revealed important new insights, pointing toward the importance of monitoring these sentinels in a rapidly changing ocean.

Cairns (1988) is often credited as first promoting seabirds as ecosystem sentinels and noted several key reasons why they were perfect for this role: (1) Seabirds are abundant, wide-ranging, and conspicuous, (2) although they feed at sea, they must return to land to nest, allowing easier observation and quantification of demographic responses, often at a fraction of the cost of traditional, ship-based oceanographic surveys, and therefore (3) parameters such as seabird reproductive success or activity budgets may respond to changing environmental conditions and provide researchers with metrics by which to assess the current state of that ecosystem.

The unprecedented 2014-2016 North Pacific marine heatwave (“the Blob”) caused extreme ecosystem disruption over an immense swath of the ocean (Cavole et al. 2016). Seabirds offered an effective and morbid indication of the scale of this disruption: Common murres (Uria aalge), an abundant and widespread fish-eating seabird, experienced widespread breeding failure across the North Pacific. Poor reproductive performance suggested that there may have been fewer small forage fish around and that these changes occurred at a large geographic scale. The Blob reached such an extreme as to kill immense numbers of adult birds, which professional and community scientists found washed up on beach-surveys; researchers estimate that an incredible 1,200,000 murres may have died from starvation during this period (Piatt et al. 2020). While the average person along the Northeast Pacific Coast during this time likely didn’t notice any dramatic difference in the ocean, seabirds were shouting at us that something was terribly wrong.

Happily, living seabirds also act as superb ecosystem sentinels. Long-term research in the Gulf of Maine by U.S. and Canadian scientists monitors the prey species provisioned by adult seabirds to their chicks. Will has spent countless hours over five summers helping to conduct this research by watching terns (Sterna spp.) and Atlantic puffins (Fratercula arctica) bring food to their young on small islands off the Maine coast. After doing this work for multiple years, it’s easy to notice that what adults feed their chicks varies from year to year. It was soon realized that these data could offer insight into oceanographic conditions and could even help managers assess the size of regional fish stocks. One of the dominant prey species in this region is Atlantic herring (Clupea harengus), which also happens to be the focus of an economically important fishery.  While the fishery targets four or five-year-old adult herring, the seabirds target smaller, younger herring. By looking at the relative amounts and sizes of young herring collected by these seabirds in the Gulf of Maine, these data can help predict herring recruitment and the relative number of adult herring that may be available to fishers several years in the future (Scopel et al. 2018).  With some continued modelling, the work that we do on a seabird colony in Maine with just a pair of binoculars can support or maybe even replace at least some of the expensive ship-based trawl surveys that are now a popular means of assessing fish stocks.

A common tern (Sterna hirundo) with a young Atlantic herring from the Gulf of Maine, ready to feed its chick (Photo courtesy of the National Audubon Society’s Seabird Institute)

For more far-ranging and inaccessible marine predators such as whales, measuring things such as dietary shifts can be more challenging than it is for seabirds. Nevertheless, whales are valuable ecosystem sentinels as well. Changes in the distribution and migration phenology of specialist foragers such as blue whales (Balaenoptera musculus) and North Atlantic right whales (Eubalaena glacialis) can indicate relative changes in the distribution and abundance of their zooplankton prey and underlying ocean conditions (Hazen et al. 2019). In the case of the critically endangered North Atlantic right whale, their recent declines in reproductive success reflect a broader regime shift in climate and ocean conditions. Reduced copepod prey has resulted in fewer foraging opportunities and changing foraging grounds, which may be insufficient for whales to obtain necessary energetic stores to support calving (Gavrilchuk et al. 2021, Meyer-Gutbrod et al. 2021). These whales assimilate and showcase the broad-scale impacts of climate change on the ecosystem they inhabit.

Blue whales that feed in the rich upwelling system off the coast of California rely on the availability of their krill prey to support the population (Croll et al. 2005). A recent study used acoustic monitoring of blue whale song to examine the timing of annual population-level transition from foraging to breeding migration compared to oceanographic variation, and found that flexibility in timing may be a key adaptation to persistence of this endangered population facing pressures of rapid environmental change (Oestreich et al. 2022). Specifically, blue whales delayed the transition from foraging to breeding migration in years of the highest and most persistent biological productivity from upwelling, and therefore listening to the vocalizations of these whales may be valuable indicator of the state of productivity in the ecosystem.

Figure reproduced from Oestreich et al. 2022, showing relationships between blue whale life-history transition and oceanographic phenology of foraging habitat. Timing of the behavioral transition from foraging to migration (day of year on the y-axis) is compared to (a) the date of upwelling onset; (b) the date of peak upwelling; and (c) total upwelling accumulated from the spring transition to the end of the upwelling season.

In a similar vein, research by the GEMM Lab on blue whale ecology in New Zealand has linked their vocalizations known as D calls to upwelling conditions, demonstrating that these calls likely reflect blue whale foraging opportunities (Barlow et al. 2021). In ongoing analyses, we are finding that these foraging-related calls were drastically reduced during marine heatwave conditions, which we know altered blue whale distribution in the region (Barlow et al. 2020). Now, for the final component of Dawn’s PhD, she is linking year-round environmental conditions to the occurrence patterns of different blue whale vocalization types, hoping to shed light on ecosystem processes by listening to the signals of these ecosystem sentinels.

A blue whale comes up for air in the South Taranaki Bight of New Zealand. photo by L. Torres.

It is important to understand the widespread implications of the rapidly warming climate and changing ocean conditions on valuable and vulnerable marine ecosystems. The cases explored here in this blog exemplify the importance of monitoring these marine megafauna sentinel species, both now and into the future, as they reflect the health of the ecosystems they inhabit.

Did you enjoy this blog? Want to learn more about marine life, research and conservation? Subscribe to our blog and get a weekly email when we make a new post! Just add your name into the subscribe box on the left panel.


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, 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.

Becker EA, Forney KA, Redfern J V., Barlow J, Jacox MG, Roberts JJ, Palacios DM (2019) Predicting cetacean abundance and distribution in a changing climate. Divers Distrib 25:626–643.

Cairns DK (1988) Seabirds as indicators of marine food supplies. Biol Oceanogr 5:261–271.

Cavole LM, Demko AM, Diner RE, Giddings A, Koester I, Pagniello CMLS, Paulsen ML, Ramirez-Valdez A, Schwenck SM, Yen NK, Zill ME, Franks PJS (2016) Biological impacts of the 2013–2015 warm-water anomaly in the northeast Pacific: Winners, losers, and the future. Oceanography 29:273–285.

Croll DA, Marinovic B, Benson S, Chavez FP, Black N, Ternullo R, Tershy BR (2005) From wind to whales: Trophic links in a coastal upwelling system. Mar Ecol Prog Ser 289:117–130.

Croll DA, Tershy BR, Hewitt RP, Demer DA, Fiedler PC, Smith SE, Armstrong W, Popp JM, Kiekhefer T, Lopez VR, Urban J, Gendron D (1998) An integrated approch to the foraging ecology of marine birds and mammals. Deep Res Part II Top Stud Oceanogr.

Gavrilchuk K, Lesage V, Fortune SME, Trites AW, Plourde S (2021) Foraging habitat of North Atlantic right whales has declined in the Gulf of St. Lawrence, Canada, and may be insufficient for successful reproduction. Endanger Species Res 44:113–136.

Hazen EL, Abrahms B, Brodie S, Carroll G, Jacox MG, Savoca MS, Scales KL, Sydeman WJ, Bograd SJ (2019) Marine top predators as climate and ecosystem sentinels. Front Ecol Environ 17:565–574.

Hazen EL, Jorgensen S, Rykaczewski RR, Bograd SJ, Foley DG, Jonsen ID, Shaffer SA, Dunne JP, Costa DP, Crowder LB, Block BA (2013) Predicted habitat shifts of Pacific top predators in a changing climate. Nat Clim Chang 3:234–238.

Hyrenbach KD, Forney KA, Dayton PK (2000) Marine protected areas and ocean basin management. Aquat Conserv Mar Freshw Ecosyst 10:437–458.

Meyer-Gutbrod EL, Greene CH, Davies KTA, Johns DG (2021) Ocean regime shift is driving collapse of the north atlantic right whale population. Oceanography 34:22–31.

Oestreich WK, Abrahms B, Mckenna MF, Goldbogen JA, Crowder LB, Ryan JP (2022) Acoustic signature reveals blue whales tune life history transitions to oceanographic conditions. Funct Ecol.

Piatt JF, Parrish JK, Renner HM, Schoen SK, Jones TT, Arimitsu ML, Kuletz KJ, Bodenstein B, Garcia-Reyes M, Duerr RS, Corcoran RM, Kaler RSA, McChesney J, Golightly RT, Coletti HA, Suryan RM, Burgess HK, Lindsey J, Lindquist K, Warzybok PM, Jahncke J, Roletto J, Sydeman WJ (2020) Extreme mortality and reproductive failure of common murres resulting from the northeast Pacific marine heatwave of 2014-2016. PLoS One 15:e0226087.

Scopel LC, Diamond AW, Kress SW, Hards AR, Shannon P (2018) Seabird diets as bioindicators of atlantic herring recruitment and stock size: A new tool for ecosystem-based fisheries management. Can J Fish Aquat Sci.

Silber GK, Lettrich MD, Thomas PO, Baker JD, Baumgartner M, Becker EA, Boveng P, Dick DM, Fiechter J, Forcada J, Forney KA, Griffis RB, Hare JA, Hobday AJ, Howell D, Laidre KL, Mantua N, Quakenbush L, Santora JA, Stafford KM, Spencer P, Stock C, Sydeman W, Van Houtan K, Waples RS (2017) Projecting marine mammal distribution in a changing climate. Front Mar Sci 4:413.

Sydeman WJ, Poloczanska E, Reed TE, Thompson SA (2015) Climate change and marine vertebrates. Science 350:772–777.