Hope lies in cooperation: the story of a happy whale!

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

I wrote my last blogpost in the midst of winter and feeling overwhelmed as I was trying to fly to the US at the peak of the omicron pandemic… Since then, morale has improved exponentially. I have spent two months in the company of my delightful GEMM lab friends, nerding over statistics, sharing scientific conversations, drinking (good!) beer and enjoying the company of this great group of people. During that stay, I was able to focus on my OPAL project more than I have ever been able to, as I set myself the goal of not getting distracted by anything else during my stay in Newport.

The only one distraction that I do not regret is a post I read one morning on the Cetal Fauna Facebook page, a group of cetacean experts and lovers who share news, opinions, photos… anything cetacean related! Someone was posting a photo of a humpback whale stranded in the 1990s’ on Coolum beach, on the east coast of Australia, which is known as a major humpback whale migratory corridor. The story said that (probably with considerable effort) the whale was refloated by many different individuals and organizations present at the beach on that day, specifically Sea World Research, Rescue & Conservation.

I felt very touched by this story and the photo that illustrated it (Figure 1). Seeing all these people come together in this risky operation to save this sea giant is quite something. And the fact that they succeeded was even more impressive! Indeed, baleen whales strand less commonly than toothed whales but their chances of survival when they do so are minimal. In addition to the actual potential damages that might have caused the whale to strand in the first place (entanglements, collisions, diseases etc.), the beaching itself is likely to hurt the animal in a permanent way as their body collapses under their own weight usually causing a cardiovascular failure (e.g., Fernández et al., 2005)⁠. The rescue of baleen whales is also simply impaired by the sheer size and weight of these animals. Compared to smaller toothed whales such as pilot whales and false killer whales that happen to strand quite frequently over some coastlines, baleen whales are almost impossible to move off the beach and getting close to them when beached can be very dangerous for responders. For these reasons, I found very few reports and publications mentioning successful rescues of beached baleen whales (e.g., Priddel and Wheeler, 1997; Neves et al., 2020).⁠

Figure 1: Stranded humpback whale on Coolum Beach, East Australia, in 1996. Look at the size of the fluke compared to the men who are trying to rescue her! Luckily, that risky operation ended well. This image won Australian Time Magazine Cover of the year. Credit: Sea World Research, Rescue and Conservation. Photo posted by P. Garbett on https://www.facebook.com/groups/CetalFauna – February 26, 2022)

Now the story gets even better… the following day I received an email from Ted Cheeseman, director and co-founder of Happywhale, a collaborative citizen science tool to share and match photographes of cetaceans (initially only humpback whales but has extended to other species) to recognize individuals based on the unique patterns of the their fluke or dorsal fin. The fluke of the whale stranded in Australia in 1991 had one and only match within the Happywhale immense dataset… and that match was to a whale seen in New Caledonia (Figure 2). “HNC338” was the one!

Figure 2: Happy whale page showing the match of HNC338 between East Australia and New Caledonia. https://happywhale.com/individual/78069;enc=284364?fbclid=IwAR1QEG_6JkpH_k2UrF-qp-9qrOboHYakKjlTj0lLbDFygjN5JugkkKVeMQw

Since I conducted my PhD on humpback whale spatial ecology in New Caledonia, I have continued working on a number of topics along with my former PhD supervisor, Dr Claire Garrigue, in New Caledonia. Although I do not remember each and every whale from her catalogue (composed of more than 1600 humpback whales as of today), I do love a good “whale tale” and I was eager to know who this HNC338 was. I quickly looked into Claire’s humpback whale database and sure enough I found it there: encountered at the end of the 2006 breeding season on September 12th, at a position of 22°26.283’S and 167°01.991’E and followed for an hour. Field notes reported a shy animal that kept the boat at a distance. But most of all, HNC338 was genetically identified as a female and was accompanied by a calf during that season! The calf was particularly big, as expected at this time of the season. What an inspiring thing to think that this whale, stranded in 1996, was resighted 10 years later in a neighboring breeding ground, apparently healthy and raising a calf of her own.

As genetic paternity analysis have been conducted on many New Caledonia calf biopsy samples as part of the Sexy Singing project conducted with our colleagues from St Andrews University in Scotland, we might be able to identify the calf’s father in this breeding stock. Thanks to the great amount of data shared and collected through Happywhale, we are discovering more and more about whale migratory patterns and behavior. It might as well be that this calf’s father was one of those whales that seem to roam over several different breeding grounds (New Caledonia and East Australia). This story is far from finished…

Figure 3: A (pretty bad!) photo of HNC338’s fluke. Luckily the Happywhale matching algorithm is very efficient and was able to detect the similarities of the fluke’s trailing edge compared to figure 1 (Cheeseman et al., 2021)⁠. Also of note, see that small dorsal fin popping out of the waters behind big mama’s fluke? That’s her calf!

From the people who pulled this whale back into the water in 1996, to the scientists and cetacean enthusiasts who shared their data and whale photos online, this story once again shows us that hope lies in cooperation! Happywhale was only created in 2015 but since then it has brought together the general public and the scientists to contribute over 465,000 photos allowing the identification of 75,000 different individuals around the globe. In New Caledonia, in Oregon and elsewhere, I hope that these collective initiatives grow more and more in the future, to the benefit of biodiversity and people.

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References

Cheeseman, T., Southerland, K., Park, J., Olio, M., Flynn, K., Calambokidis, J., et al. (2021). Advanced image recognition: a fully automated, high-accuracy photo-identification matching system for humpback whales. Mamm. Biol. doi:10.1007/s42991-021-00180-9.

Fernández, A., Edwards, J. F., Rodríguez, F., Espinosa De Los Monteros, A., Herráez, P., Castro, P., et al. (2005). “Gas and fat embolic syndrome” involving a mass stranding of beaked whales (Family Ziphiidae) exposed to anthropogenic sonar signals. Vet. Pathol. 42, 446–457. doi:10.1354/vp.42-4-446.

Neves, M. C., Neto, H. G., Cypriano-Souza, A. L., da Silva, B. M. G., de Souza, S. P., Marcondes, M. C. C., et al. (2020). Humpback whale (megaptera novaeangliae) resighted eight years after stranding. Aquat. Mamm. 46, 483–487. doi:10.1578/AM.46.5.2020.483.

Priddel, D., and Wheeler, R. (1997). Rescue of a Bryde’s whale Balaenoptera edeni entrapped in the Manning River, New South Wales: Unmitigated success or unwarranted intervention? Aust. Zool. 30, 261–271. doi:10.7882/AZ.1997.002.

Cross-taxa collaborations: a look at the value of human and cetacean partnerships.

Imogen Lucciano, Graduate student, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab.

For marine science to be successful and impactful, it is crucial for collected data and results of analyses to be shared as widely as possible. This sharing should occur with the research community itself (which of course saves time and helps ignite the big, impactful ideas), and also amongst the public, in government, the fishing industry, big energy businesses, the military, and shipping industries as well. All these entities can relate in some way to the use of the oceans. Our increased collective knowledge can help us make conscious and intelligent management choices that will promote healthy oceans and in turn provide more resources to humans as well.

Though I am only just breaking the ice in my marine science education, I am already experiencing my first tastes of what this collaboration can look like. My graduate thesis focuses on the acoustic and observational detections of fin whales, an endangered species, as they relate to environmental characteristics in the NE Pacific. I am still in the early stages collecting data with the HALO project, but for now it is important to get started reviewing what’s currently available in the field. GEMM lab’s OPAL project, led by Dr. Leigh Torres and Dr. Solene Derville, was quick to provide me with their fin whale sightings data collected over the past few years, as well as share some of their great fin whale photos (Fig. 1). Clearly, I am already becoming rich through this association.

Figure 1. Two fin whales surface off the Oregon coast. Photographed by Leigh Torres during an OPAL helicopter survey in September 2021 under NMFS permit # 21678.

My career interests revolve around filling knowledge gaps of cetacean behaviors, so I often find myself associating what’s happening in my life to what I am reading currently as it relates to this field of research. My most recent blog, highlighted my need to relax occasionally with play and prompted me to consider how play is defined in cetacean behavior. So, with the ignition of my graduate research and this first aforementioned taste of scientific collaboration, I synaptically thought about a recent study of interspecies collaborative hunting between dolphins and humans that was co-authored by the Marine Mammal Institute’s Dr. Mauricio Cantor. Here, bottlenose dolphins who have learned to herd fish to shore, stick together and use their skills to move schools of fish toward local fishermen standing by with nets. The dolphins then provide a signal to the fishermen, the nets are cast at just the right time, and the dolphins forage on the fish trapped between the fishermen and the nets (Daura-Jorge, Cantor, et al., 2012). Both the dolphins and the fishermen greatly benefit by working together. I found this study thought-provoking; I have not seen anything quite like this interspecies association.

National Geographic video provides close perspective of the Laguna, Brazil fishermen working together with dolphins to net fish. https://www.youtube.com/watch?v=8kMGJ8T3-Pg.

In the interest of potentially finding more cross-taxa cetacean relationships, I dug into the literature and found a few more interspecies associations to note. The first article that took me aback was a 2017 report detailing humpback whales defending other marine mammal species by interfering with the hunting practices of transient killer whales (Pitman et al., 2017). Killer whales are apex predators who hunt marine mammals, to include pinnipeds, adult baleen whales and often the calves of baleen whales. Slow, rotund baleen whales (right whales, gray whales, and humpbacks) are known to use their immense size and large appendages to fight off killer whales. What is unique with this study is that humpback whales were observed not only protecting their own calves from predation but also using a mobbing tactic to protect other cetacean species (minke whales, gray whales, Dall’s porpoises, and others) and pinnipeds (Steller sea lions, California sea lions, Weddell seals, and others; Fig. 2) as well, showing acts of potential altruism in cetaceans (Pitman et al., 2017).

Figure 2. Humpback whale moving in to interfere with a killer whale hunting a seal. Photo credit: Robert Pitman, https://whalescientists.com/humpback-whales-altruism/.

The next interspecies association catching my eye came from studies detailing the two largest marine mammals, blue and fin whales, reproducing together. Though the two species are relatively alike in having large sleek physiques, they are very different in their known migratory and acoustic behaviors, so it doesn’t seem obvious or likely the two would mate. However, following the genetic testing of a whale near Iceland that displayed an unusual phenotype, researchers were able to determine that the whale did in fact contain the DNA of both species (Pampoulie et al., 2020). These blue/fin hybrids have been spotted in several locations worldwide and they are even found to be fertile. A recent study of a successfully tagged and observed blue/fin hybrid called, “Flue” (Fig. 3), co-authored by Dr. Daniel Palacios of MMI’s WHET Lab, found that though the animal possessed a phenotype mostly descriptive of fin whale, Flue appeared to follow blue whale migratory behavior (moving farther north along the California coast to forage in the summer and then moving to southern breeding ground waters along the coast of Mexico). These researchers suggest that blue/fin hybrid whales are common and postulate whether these animals are the source of an unmatched 52 Hz whale call sometimes recorded in the North Pacific (Jefferson et al., 2021).

Figure 3. Highly observed and documented blue/fin whale hybrid, called “Flue”, spotted off the coast of Santa Barbara, CA, USA. Photo credit: Adam Ernster, Condor Express Media, https://www.youtube.com/watch?v=4LjH2-naRPE&feature=youtu.be&app=desktop.

Lastly (and perhaps my favorite of the papers of the collection), there is a report published in 2019 detailing a closely followed bottlenose dolphin female who adopted a young melon-headed whale calf near French Polynesia in the South Pacific (Fig. 4). Though cetaceans have been known to participate in allonursing, a form of alloparental care in which adult females will nurse another’s offspring of the same species, an interspecific adoption has rarely been reported. This mother-calf interspecies pair were observed together just after the adoptive mother gave birth to another calf, so it was impossible that the adopted calf was a potential hybrid. Furthermore, the two species have overlapping populations in this area of the South Pacific and thus it was concluded that the female dolphin had accepted a lost calf as her own (Carzon et al., 2019). Lactation is energetically costly, and considering the dolphin already had another calf to feed, the fact that she accepted the adopted calf, was observed nursing it, and developed a lengthy bond with it is remarkable.

Figure 4. Bottlenose dolphin female with her adopted melon-headed whale calf near French Polynesia in the South Pacific (Carzon et al., 2019).

I admit it was more fun than work to dig into these interspecies associations this week, because they depict how rich our world can be when animals (including humans) evoke positive associations across taxa. Reverting into my fin whale research, I cannot wait to see where my analysis will lead. I am eager to share my results, begin collaborations with other researchers and eventually present it to the public with the hopes of developing positive associations between humans and the marine world.

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Literature Cited

Carzon, P., Delfour, F., Dudzinski, K. et al. 2019. Cross-genus adoption in delphinids: One example with taxonomic discussion. Ethology: Behavioral Notes, 125: 669-676.

Daura-Jorge, F., Cantor, M., Ingram, S. et al. 2012. The structure of a bottlenose dolphin society is coupled to a unique foraging cooperation with artisanal fisherman. Biology Letters, 8: 702-705.

Jefferson, T., Palacios, D., Calambokidis, J. et al. 2021. Sightings and satellite tracking of a blue/fin whale hybrid in its wintering and summering ranges in the eastern north pacific. Advances in Oceanography & Marine Biology, 2 (4). http://dx.doi.org/10.33552/AOMB.2021.02.000545.  

Pampoulie, C., Gislason, D., Olafsdottir, G. et al. 2020. Evidence of unidirectional hybridization and second-generation adult hybrid between the two largest animals on Earth, the fin and blue whales. Evolutionary Applications, 14: 314-321.

Pitman, R., Deecke, V., Gabriele, C., et al. 2016. Humpback whales interfering when mammal-eating killer whales attack other species: Mobbing behavior and interspecific altruism? Marine Mammal Science, 33 (1): 7-58. https://doi.org/10.1111/mms.12343.

Back to the Future: The return of scientific conferences

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

The pandemic has taught me that certain skills – including ones I never recognized as such – can atrophy. How do I construct an outfit that involves actual pants instead of gym shorts? How do I make a lunch that is portable and can be eaten outside my home?

These are things that I’ve had to relearn over the last year, as I increasingly leave my virtual work world and move back into the physical world. Recently, the new ways in which the world is opening up again have pushed me to brush off another skill – how do I talk to other people about my work?

The pandemic has necessarily made the world a bit more insular. A year and a half into my graduate career, I’ve mostly discussed my work within the cozy cocoon of my lab groups and cohort. In particular, I’ve lived the last few months in that realm of research that is so specific and internal that almost no one else fully understands or cares about what I’m doing: I’ve spent days tangled up in oodles of models, been woken up at night by dreams about coding, and sweated over the decimal points of statistical deviance-explained values. 

This period of scientific navel gazing abruptly ended this February. In the space of ten days, I presented at my first in-person conference during graduate school, gave a short talk at my first international conference, and gave my longest talk yet to a public audience. After reveling in the minutiae of research for months, it was so valuable to be forced to take a step back, think about the overarching narrative of this work, and practice telling that story to different audiences. 

A February talk for the Oregon chapter of the American Cetacean Society gave me the chance to tell the story of my research to a broad audience.

Presenting this work to an in-person audience for the first time was especially rewarding. With a physical (!) poster in hand, I headed out to Newport for the annual meeting of the Oregon Chapter of The Wildlife Society. The GEMM Lab really took this conference by storm – Leigh gave a plenary talk on the meeting’s theme of “Dynamic Oceans, Shifting Landscapes”, Lisa chaired a session and gave a talk about trophic relationships between kelp and whales, and Miranda presented a poster on the new Holistic Assessment of Living marine resources off the Oregon coast (HALO) project.

This great GEMM Lab presence gave me the opportunity to reference everyone else’s work as I shared my own, and to think about the body of work we do as a group and the coherence in research themes that different projects share. I almost lost my voice by talking for the entire duration of the poster session, and was energized by the opportunity to share this work with so many interested people.

The GEMM Lab and other OSU Marine Mammal Institute members presented alongside terrestrial researchers on the theme of “Dynamic Oceans, Shifting Landscapes”.

Just a few days later, the biennial Ocean Sciences Meeting began. Dawn presented on forecasting the distribution of blue whales in New Zealand’s South Taranaki Bight region, and several members of the Krill Seeker Lab, led by my co-advisor Dr. Kim Bernard, presented their own zooplankton ecology research.

Originally scheduled for Hawaii, this meeting was instead held virtually as a safety precaution against Covid-19. Nevertheless, the diversity of talks and time spent gathering online still gave me the sense of being part of an international ocean science community. People attended from every time zone, and watching early-morning talks while wearing pajamas with Solene, Dawn, and Quin the dog is officially one of my new favorite conference experiences.

In addition to the chance to discuss science with other students and researchers, it was great to have the opportunity to step back from our normal routines a bit. The Krill Seeker Lab did the conference-organized 5K walk together (in intermittent rain, of course) and our team even came within one point of winning the trivia contest. All the while, we were hopping in and out of poster sessions and talks, realizing that virtual conferences can be just as busy as in-person ones.

Taking a 5k-long break from watching talks! From left to right: Rachel Kaplan, Kim Bernard, Giulia Wood, and Kirsten Steinke.

Over the last two years, one of the things the pandemic has made me appreciate the most is the ability to gather. Dinner with friends, holidays with family – the ability to be together is far more tentative and precious than I realized during the “before times.” Now, as we start tiptoeing back into normal life a bit more, I’m appreciating the ability to gather for science and looking forward to more conferences in the future.

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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.

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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, 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.

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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.

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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.

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