As a newly-minted PhD student, starting graduate school has so far been everything I dreamt — and a bit more. I expected the excitement of meeting my cohort and professors, and starting classes. The apocalyptic drive to campus through a fiery sky as fires burned across Oregon, and the week after spent solely indoors, I did not.
As I’ve settled into Corvallis, my program, and navigating the roadblocks 2020 keeps throwing our way, I have been so grateful for the warm (virtual) welcome by my lab groups, professors, and fellow students. One of the most impressive displays of flexibility and adaptability so far is the ever-evolving field course I am currently taking.
Called “Cascadia,” this course provides an introduction to the range of geological, physical, ecological, and biogeochemical topics that exist within the Pacific Northwest, and explores the linkages between these areas. The course’s goal is to introduce incoming CEOAS (College of Earth, Ocean, and Atmospheric Sciences) students to the surrounding landscape, and to the ways that human systems interact with that landscape.
The professors teaching Cascadia — Drs. Frederick Colwell, Emily Shroyer, and George Waldbusser — have done an amazing job adapting the course to unprecedented circumstances. Over the summer, safety measures due to the pandemic required them to move the course to a largely online format, with only three planned day trips (typically the course is a full ten-day road trip around the state). Over the last week, the fires raging around Oregon have forced them to adapt the course repeatedly in real time, postponing field trips based on air quality forecasts and site closures.
During a typical year in the Cascadia course, the incoming students learn while exploring, camping, and hiking their way around a number of sites around Oregon. This year, our classmates are scattered around the country and our explorations have taken place in a Zoom room — but that hasn’t stopped the experience from being great.
Several professors shared their expertise with us through a series of talks that covered the ecology and history of the Willamette River, Pacific Northwest volcanoes, tsunami safety and preparation, and even wildfire ecology. In addition to talks by subject matter experts, each student delved into and presented on a topic of their choice, allowing us to learn from one another about everything from edible plants, to Oregon craft beers, to human movements throughout the Willamette River valley. We also enjoyed gorgeous pictures of Oregon’s mountains, coast, and desert, and received recommendations for trips and hikes that everyone is excited to explore.
As of the time of writing this blog, I’m excited to say that things may look a little different tomorrow — rain and improved air quality are in the forecast, and the Cascadia crew is planning to venture out to the coast for our first field trip! We’ll be learning on-site about the Oregon Coast Range and coastal dynamics, climate, and processes. This will actually be my first time on the Oregon coast, but definitely not my last.
A big part of my excitement about this research project lies in the way it intersects natural and human systems, just as we have been exploring through the Cascadia course. I am interested in how marine mammal distribution and behavior intersect with human systems — and how understanding these interactions can inform management and conservation efforts. I am thrilled to be a new member of the GEMM Lab, and to be starting (remote) classes and this research. For now, I’m wishing everyone good air quality and a safe fall!
In the vast and dynamic marine environment, food is notoriously patchy and ephemeral . Predators such as marine mammals and seabirds must make a living in this dynamic environment by locating and capturing those prey patches. Baleen whales such as blue and humpback whales have a feeding strategy called “lunge feeding”, whereby they accelerate forward and open their massive jaws, engulf prey-laden water in their buccal pouch that expands like an accordion, and filter the water out through baleen plates so that they are left with a mouthful of food (Fig. 1) . This approach is only efficient if whales can locate and target dense prey patches that compensate for the energetic costs of diving and lunging . Therefore, not only do these large predators need to locate enough food to survive in the expansive and ever-changing ocean, they need to locate food that is dense enough to feed on, otherwise they actually lose more energy by lunging than they gain from the prey they engulf.
Why do baleen whales rely on such a costly feeding approach? Interestingly, this tactic emerged after the evolution of schooling behavior of prey such as zooplankton and forage fish (e.g., herring, anchovy, sand lance) . Only because the prey aggregate in dense patches can these large predators take advantage of them by lunge feeding, and by engulfing a whole large patch they efficiently exploit these prey patches. Off the coast of California, where krill aggregations are denser in deeper water, blue whales regularly dive to depths of 100-300 m in order to access the densest krill patches and get the most bang for their buck with every lunge . In New Zealand, we have found that blue whales exploit the dense krill patches near the surface to maximize their energetic gain , and have documented a blue whale bypassing smaller krill patches that presumably were not worth the effort to feed on.
By now hopefully I have convinced you of the importance of dense prey patches to large whales looking for a meal. It is not necessarily only a matter of total prey biomass in an area that is important to a whale, it is whether that prey biomass is densely aggregated. What makes for a dense prey patch? Recent work has shown that forage species, namely krill and anchovies, swarm in response to coastal upwelling . While upwelling events do not necessarily change the total biomass of prey available to a whale over a spatial area, they may aggregate prey to a critical density to where feeding by predators becomes worthwhile. Forage species like zooplankton and small fish may school because of enhanced food resources, for predator avoidance, or reproductive grouping. While the exact behavioral reason for the aggregation of prey may still only be partially understood, the existence of these dense patches allows the largest animals on the planet to survive.
Another big question is, how do whales actually find their food? In the vast, seemingly featureless, and ever-changing ocean environment, how does a whale know where to find a meal, and how do they know it will be worthwhile before they take a lunge? In a review paper written by GEMM Lab PI Dr. Leigh Torres, she suggests it is all a matter of scale . On a very large scale, baleen whales likely rely on oceanographic stimuli to home in on areas where prey are more likely to be found. Additionally, recent work has demonstrated that migrating blue whales return to areas where foraging conditions were best in previous years, indicating a reliance on memory [9,10]. On a very fine scale, visual cues may inform how a blue whale chooses to lunge [6,8,11].
What does it matter what a blue whale’s favorite type of meal is? Besides my interest in foundational research in ecology such as predator-prey dynamics, these questions are fundamental to developing effective management approaches for reducing impacts of human activities on whales. In the first chapter of my PhD, I examined how oceanographic features of the water column structure krill aggregations, and how blue whale distribution is influenced by oceanography and krill availability . Currently, I am deep into my second chapter, analyzing the pathway from wind to upwelling to krill to blue whales in order to better understand the links and time lags between each step. Understanding the time lags will allow us to make more informed models to forecast blue whale distribution in my third chapter. Environmental managers in New Zealand plan to establish a protected area to conserve the population of blue whales that I study  on their foraging grounds. Understanding where blue whales will be distributed, and consequently how their distribution patterns might shift with environmental conditions or overlap with human activities, comes down the fundamental question I started this blog post with: What makes a good meal for a hungry whale?
1. Hyrenbach KD, Forney KA, Dayton PK. 2000 Marine protected areas and ocean basin management. Aquat. Conserv. Mar. Freshw. Ecosyst.10, 437–458. (doi:10.1002/1099-0755(200011/12)10:6<437::AID-AQC425>3.0.CO;2-Q)
2. Goldbogen JA, Cade DE, Calambokidis J, Friedlaender AS, Potvin J, Segre PS, Werth AJ. 2017 How Baleen Whales Feed: The Biomechanics of Engulfment and Filtration. Ann. Rev. Mar. Sci.9, 367–386. (doi:10.1146/annurev-marine-122414-033905)
3. Goldbogen JA, Calambokidis J, Oleson E, Potvin J, Pyenson ND, Schorr G, Shadwick RE. 2011 Mechanics, hydrodynamics and energetics of blue whale lunge feeding: efficiency dependence on krill density. J. Exp. Biol.214, 131–146. (doi:10.1242/jeb.048157)
4. Cade DE, Carey N, Domenici P, Potvin J, Goldbogen JA. 2020 Predator-informed looming stimulus experiments reveal how large filter feeding whales capture highly maneuverable forage fish. Proc. Natl. Acad. Sci. U. S. A. (doi:10.1073/pnas.1911099116)
5. Hazen EL, Friedlaender AS, Goldbogen JA. 2015 Blue whales (Balaenoptera musculus) optimize foraging efficiency by balancing oxygen use and energy gain as a function of prey density. Sci. Adv.1, e1500469–e1500469. (doi:10.1126/sciadv.1500469)
6. 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 (doi:10.7717/peerj.8906)
7. Benoit-Bird KJ, Waluk CM, Ryan JP. 2019 Forage Species Swarm in Response to Coastal Upwelling. Geophys. Res. Lett.46, 1537–1546. (doi:10.1029/2018GL081603)
8. Torres LG. 2017 A sense of scale: Foraging cetaceans’ use of scale-dependent multimodal sensory systems. Mar. Mammal Sci.33, 1170–1193. (doi:10.1111/mms.12426)
9. Abrahms B et al. 2019 Memory and resource tracking drive blue whale migrations. Proc. Natl. Acad. Sci. U. S. A. (doi:10.1073/pnas.1819031116)
10. Szesciorka AR, Ballance LT, Širovi A, Rice A, Ohman MD, Hildebrand JA, Franks PJS. 2020 Timing is everything: Drivers of interannual variability in blue whale migration. Sci. Rep.10, 1–9. (doi:10.1038/s41598-020-64855-y)
11. Friedlaender AS, Herbert-Read JE, Hazen EL, Cade DE, Calambokidis J, Southall BL, Stimpert AK, Goldbogen JA. 2017 Context-dependent lateralized feeding strategies in blue whales. Curr. Biol. (doi:10.1016/j.cub.2017.10.023)
12. 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. (doi:https://doi.org/10.3354/meps13339)
13. Barlow DR et al. 2018 Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger. Species Res.36, 27–40. (doi:https://doi.org/10.3354/esr00891)
Recently, I had the opportunity to attend the International Statistical Ecology Conference (ISEC), a biennial meeting of researchers at the interface of ecology and statistics. I am a marine ecologist, fascinated by the interactions between animals and the dynamic ocean environment they inhabit. If you had asked me five years ago whether I thought I would ever consider myself a statistician or a computer programmer, my answer would certainly have been “no”. Now, I find myself studying the ecology of blue whales in New Zealand using a variety of data streams and methodologies, but a central theme for my dissertation is species distribution modeling. Species distribution models (SDMs) are mathematical algorithms that correlate observations of a species with environmental conditions at their observed locations to gain ecological insight and predict spatial distributions of the species (Fig. 1; Elith and Leathwick 2009). I still can’t say I would identify as a statistician, but I have a growing appreciation for the role of statistics to gain inference in ecology.
Before I continue, let’s take a look at just a few definitions from Merriam-Webster’s dictionary:
Statistics: a branch of mathematics dealing with the collection, analysis, interpretation, and presentation of masses of numerical data
Ecology: a branch of science concerned with the interrelationship of organisms and their environments
Inference: a conclusion or opinion that is formed because of known facts or evidence
Ecological data are notoriously noisy, messy, and complex. Statistical tests are meant to help us understand whether a pattern in the data is different from what we would expect through random chance. When we study how organisms interact with one another and their environment, it is impossible to completely capture all elements of the ecosystem. Therefore, ecology is a field ripe with challenges for statisticians. How do we quantify a meaningful biological signal amidst all the noise? How can we gain inference from ecological data to enhance knowledge, and how can we use that knowledge to make informed predictions? Marine mammals are notoriously difficult to study. They inhabit an environment that is relatively inaccessible and inhospitable to humans, they occur in low numbers, they are highly mobile, and they are rarely visible. All ecological data are difficult and noisy and riddled with small sample sizes, but counting trees presents fewer logistical challenges than counting moving whales in an ever-changing open-ocean setting. Therefore, new methodologies in areas like species distribution modeling are often developed using large, terrestrial datasets and eventually migrate to applications in the marine environment (Robinson et al. 2011).
Many presentations I attended at the conference were geared toward moving beyond correlative SDMs. SDMs were developed to correlate species occurrence patterns with features of the environment they inhabit (e.g. temperature, precipitation, terrain, etc.). However, those relationships do not actually explain the underlying mechanism of why a species is more likely to occur in one environment compared to another. Therefore, ecological statisticians are now using additional information and modeling approaches within SDMs to incorporate information such as species co-occurrence patterns, population demographic information, and physiological constraints. Building SDMs to include such process-explicit information allows us to make steps toward understanding not just when and where a species occurs, but why.
Machine learning is an area that continues to advance and open doors to new applications in ecology. Machine learning approaches differ fundamentally from classical statistics. In statistics, we formulate a hypothesis, select the appropriate model to test that hypothesis (for example, linear regression), then test how well the data fit the model (“Is the relationship linear?”), and test the strength of that inference (“Is the linear pattern different from what we would expect due to random chance?”). Machine learning, on the other hand, does not use a predetermined notion of relationships between variables. Rather, it tries to create an algorithm that fits the patterns in the data. Statistics asks how well the data fit a model, and machine learning asks how well a model fits the data.
Machine learning approaches allow for very complex relationships to be included in models and can be excellent for making predictions. However, sometimes the relationships fitted by a machine learning algorithm are so complex that it is not possible to infer any ecological meaning from them. As one ISEC presenter put it, in machine learning “the computer learns but the scientist does not”. The most important thing when selecting your methodology is to remember your question and your goal. Do you want to understand the mechanism of why an animal is where it is? Or do you not need to understand the driver, but rather want to make the best predictions of where an animal will be? In my case, the answer to that question differs from one of my PhD chapters to the next. We want to understand the functional relationships between oceanography, krill availability, and blue whale distribution (Barlow et al. 2020), and subsequently we want to develop forecasting models that can reliably predict blue whale distribution to inform conservation efforts (Fig. 2).
ISEC was an excellent opportunity for me to break out of my usual marine mammal-centered bubble and get a taste of what is happening on the leading edge of statistical ecology. I learned about the latest approaches and innovations in species distribution modeling, and in the process I also learned about trees, koalas, birds, and many other organisms from around the world. A fun bonus of attending a methods-focused conference is learning about completely new study species and systems. There are many ways of approaching an ecological question, gaining inference, and making predictions. I look forward to incorporating the knowledge I gained through ISEC into my own research, both in my doctoral work and in applications of new methods to future research projects.
Barlow, D.R., Bernard, K.S., Escobar-Flores, P., Palacios, D.M., and Torres, L.G. 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. doi:https://doi.org/10.3354/meps13339.
Elith, J., and Leathwick, J.R. 2009. Species Distribution Models: Ecological Explanation and Prediction Across Space and Time. Annu. Rev. Ecol. Evol. Syst. 40(1): 677–697. doi:10.1146/annurev.ecolsys.110308.120159.
Robinson, L.M., Elith, J., Hobday, A.J., Pearson, R.G., Kendall, B.E., Possingham, H.P., and Richardson, A.J. 2011. Pushing the limits in marine species distribution modelling: Lessons from the land present challenges and opportunities. doi:10.1111/j.1466-8238.2010.00636.x.
For my PhD, I am using a variety of data sources and analytical tools to study the ecology and distribution of blue whales in New Zealand. I live on the Oregon Coast, across the world and in another season from the whales I study. I love where I live and I am passionate about my work, but I do sometimes feel removed from the whales and the ecosystem that are the focus of my research.
Recently, I have turned my attention to processing acoustic data recorded in our study region in New Zealand between 2016 and 2018. In the fall, I developed detector algorithms to identify possible blue whale vocalizations in the recording period, and now I am going through each of the detections to validate whether it is indeed a blue whale call or not. Looking closely at spectrograms for hours and hours is a change of pace from the analysis and writing I have been doing recently. Namely, I am looking at biological signals – not lines of code and numbers on a screen, but depictions of sounds that blue whales produced. I have to say, it is the “closest” I have felt to these whales in a long time. Scrolling through thousands of spectrograms of blue whale calls leaves room for my mind to wander, and I recently had the realization that those whales have absolutely no idea that on the other side of the Pacific Ocean, there are a few scientists dedicating years of their lives to understand and protect them. Which led me to another realization: we know so much more about blue whales in New Zealand now than we did 10 years ago. In fact, we know so much more than we did even a year ago.
Nine years ago, Dr. Leigh Torres had a cup of coffee with a colleague who recounted observer reports of several blue whales during a seismic survey of the South Taranaki Bight region (STB) of New Zealand. This conversation sparked her curiosity, and led to the formulation of a hypothesis that the STB was in fact an unrecognized feeding ground for blue whales in the southern hemisphere (Torres 2013).
After three field seasons and several years of dedicated work, the hypothesis that the STB region is important for blue whales was validated. By drawing together multiple data streams and lines of evidence, we now know that New Zealand is home to a unique population of blue whales, which are genetically distinct from all other documented populations in the Southern Hemisphere. Furthermore, they use the STB for multiple critical life history functions such as feeding, nursing and calf raising, and they are present there year-round (Barlow et al. 2018).
Once we documented the New Zealand population, we were left with perhaps even more questions than we started with. Where do they feed, and why? Are they feeding and breeding there? Does their distribution change seasonally? What is the health of the population? Are they being impacted by industrial activity and human impacts such as noise in the region? We certainly do not have all the answers, but we have been piecing together an increasingly comprehensive story about these whales and their ecology.
For example, we now know that blue whales in New Zealand average around 19 meters in length, which we calculated by measuring images taken via drones and using an analysis program developed in the GEMM Lab (Burnett et al. 2018). The use of drones has opened up a whole new world for studying health and behavior in whales, and we recently used video footage to better understand the movement and kinematics of how blue whales engulf their krill prey. Furthermore, we know that blue whales may preferentially feed on dense krill aggregations at the surface, and that this surface feeding strategy may be an energetically favorable strategy in this part of the world (Torres et al. 2020).
We have also assessed one aspect of the health of blue whales by describing their skin condition. By analyzing thousands of photographs, we now know that nearly all blue whales in New Zealand bear the scars of cookie cutter shark bites, which they seem more likely to acquire at more northerly latitudes, and that 80% are affected by blister lesions (Barlow et al. 2019). Next, we are beginning to draw together multiple data streams such as body condition and hormone analysis, paired with skin condition, to form a detailed understanding about the health of this population.
Most recently we have produced a study describing how oceanography, prey and blue whales are connected within this region of New Zealand. The STB region is home to a wind-driven upwelling system that drives productivity and leads to aggregations of krill, which in turn provide sustenance for blue whales to feed on. By compiling data on oceanography and water column structure, krill availability, and blue whale distribution, we now have a solid understanding of this trophic pathway: how oceanography structures prey, and how blue whales respond to both prey and oceanography (Barlow et al. 2020). Furthermore, we are beginning to understand how those relationships may look under changing ocean conditions, with global sea temperatures rising and the increasing frequency and intensity of marine heatwaves.
The knowledge we have accumulated better enables managers to make informed decisions for the conservation of these blue whales and the ecosystem they inhabit. To me, there are several take-away messages from the story that continues to unfold about these blue whales. One is the importance of following a hunch, and then gathering the necessary tools and team to explore and tackle challenging questions. An idea that started over a cup of coffee and many years of hard work and dedication have led to a whole new body of knowledge. Another message is that the more questions you ask and the more questions you try to answer, the more questions you are often left with. That is a beautiful truth about scientific inquiry – the questions we ask drive the knowledge we uncover, but that process is never complete because new questions continue to emerge. Finally, it is easy to get swept up in details, outputs, timelines, and minutia, and every now and then it is important to take a step back. I have appreciated taking a step back and musing on the state of our knowledge about these whales, about how much we have learned in less than 10 years, and mostly about how many answers and new questions are still waiting to be uncovered.
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.
Barlow DR, Pepper AL, Torres LG (2019) Skin Deep: An Assessment of New Zealand Blue Whale Skin Condition. Front Mar Sci.
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.
Burnett JD, Lemos L, Barlow DR, Wing MG, Chandler TE, Torres LG (2018) Estimating morphometric attributes on baleen whales using small UAS photogrammetry: A case study with blue and gray whales. Mar Mammal Sci.
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.
By: Alexa Kownacki, Ph.D. Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab
While sitting on my porch and watching the bees pollinate the blooming spring flowers, I intermittently pause to scratch the hives along my shoulders and chest. In the middle of my many Zoom calls, I mute myself and stop my video because a wave of pollen hits my face and I immediately have to sneeze. With this, I’m reminded: Welcome to prime allergy season in the Northern Hemisphere. As I was scratching my chronic idiopathic urticaria (hives caused by an overactive immune system), I asked myself “Do dolphins get hives?” I had no idea. I know most terrestrial mammals can and do—just yesterday, one of the horses in the nearby pasture was suffering from a flare of hives. But, what about aquatic and marine mammals?
As with most research on marine mammal health, knowledge is scare and is frequently limited to studies conducted on captive and stranded animals. Additionally, most of the current theories on allergic reactions in marine mammals are based on studies from terrestrial wildlife and humans. Because nearly all research on histamine pathways centers on terrestrial animals, I wanted to see what information exists the presence of skin allergies in marine mammals.
Allergic reactions trigger a cascade within the body, beginning with the introduction of a foreign body, which for many people is pollen. The allergen binds to antibodies that are produced to fight potentially harmful substances. Once this allergen binds to different types of cells, including mast cells, chemicals like histamines are released. Histamines cause the production of mucus and constriction of blood vessels, and thus are the reason your eyes water, your nose runs, or you start coughing.
As you probably can tell just by looking at a marine mammal, they have thicker skin and fewer mucus membranes that humans, due to the fact that they live in the water. However, mast cells or mast cell-like cells have been described in most vertebrate lineages including mammals, birds, reptiles, amphibians, and bony fishes (Hellman et al. 2017, Reite and Evenson 2006). Mast cell-like cells have also been described in an early ancestor of the vertebrates, the tunicate, or sea squirt (Wong et al. 2014). Therefore, allergic-reaction cascades that may present as hives, red and itchy eyes or nose in humans, also exist in marine mammals, but perhaps cause different or less visible symptoms.
Skin conditions in cetaceans are gathering interest within the marine mammal health community. Even our very own Dawn Barlow, Dr. Leigh Torres, and Acacia Pepperassessed the skin conditions in New Zealand blue whales in their recent publication. Most visible skin lesions or markings on cetaceans are caused by parasites, shark bits, fungal infections, and fishery or boat interactions (Leone et al. 2019, Sweeney and Ridgway 1985). However, there is very little scientific literature about allergic reactions in marine mammals, let alone cetaceans. That being said, I managed to find a few critical pieces of information supporting the theory that marine mammals do in fact have allergies that can produce dermal reactions similar to hives in humans.
In one study, three captive bottlenose dolphins developed reddened skin, sloughing, macules, and wheals on their ventral surfaces (Monreal-Pawlowsky et al. 2017). The medical staff first noticed this atopic dermatitis in 2005 and observed the process escalate over the next decade. Small biopsy samples from the affected areas on the three dolphins coincided with the appearance of four pollens in the air within the geographic region: Betula, Pistacia, Celtis, and Fagus (Monreal-Pawlowsky et al. 2017). Topical prednisone treatments were applied to the affected areas at various dosages that slowly resolved the skin irritations. Researchers manufactured an allergy vaccine using a combination of the four pollens in hopes that it would prevent further seasonal outbreaks, but it was unsuccessful. In the coming years, the facility intends to adjust the dosages to create a successful vaccine.
In addition to the above study, there is an unpublished case of suspected allergic reaction to another pollen that produces a pruritic reaction on the ventral areas of dolphins on a seasonal basis (Vicente Arribes, personal communication). Although there are only a few documented cases of environmentally-triggered allergic reactions that are visible on the dermal layer of cetaceans, I believe this evidence makes the case that some cetaceans suffer from allergies much like us. So, next time you’re enjoying the beautiful blooms and annoyingly scratch your eyes, know that you are not alone.
Barlow DR, Pepper AL and Torres LG (2019) Skin Deep: An Assessment of New Zealand Blue Whale Skin Condition. Front. Mar. Sci. 6:757.doi: 10.3389/fmars.2019.00757
Hellman LT, Akula S, Thorpe M and Fu Z (2017) Tracing the Origins of IgE, Mast Cells, and Allergies by Studies of Wild Animals. Front. Immunol. 8:1749. doi: 10.3389/fimmu.2017.01749
Leone AB, Bonanno Ferraro G, Boitani L, Blasi MF. Skin marks in bottlenose dolphins (Tursiops truncatus) interacting with artisanal fishery in the central Mediterranean Sea. PLoS One. 2019;14(2):e0211767. Published 2019 Feb 5. doi:10.1371/journal.pone.0211767
Monreal-Pawlowsky T, Fernández-Bellon H, Puigdemont A (2017) Suspected Allergic Reaction in Bottlenose Dolphins (Tursiops truncatus). J Vet Sci Ani Husb 5(1): 108. doi: 10.15744/2348-9790.5.108
Reite OB, Evensen O. Inflammatory cells of teleostean fish: a review focusing on mast cells/eosinophilic granule cells and rodlet cells. Fish Shellfish Immunol (2006) 20:192–208. doi:10.1016/j.fsi.2005.01.012
Sweeney, J. C., & Ridgway, S. H. (1975). Common diseases of small cetaceans. J. Am. Vet. Med. Assoc, 167(7), 533-540.
Wong GW, Zhuo L, Kimata K, Lam BK, Satoh N, Stevens RL. Ancient originof mast cells. Biochem Biophys Res Commun (2014) 451:314–8. doi:10.1016/j.bbrc.2014.07.124
Clara Bird, Masters Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab
Happy new year from the GEMM lab! Starting graduate school comes with a lot of learning. From skills, to learning about how much there is to learn, to learning about the system I will be studying in depth for the next few years. This last category has been the most exciting to me because digging into the literature on a system or a species always leads to the unearthing of some fascinating and surprising facts. So, for this blog I will write about one of the aspects of gray whale foraging that intrigues me most: benthic feeding and its impacts.
How do gray whales
Gray whales are a unique species. Unlike other baleen whales, such as humpback and blue whales, gray whales regularly feed off the bottom of the ocean (Nerini, 1984). They roll to one side and swim along the bottom, they then suction up (by depressing their tongue) the sediment and prey, then the sediment and water is filtered out of the baleen. In fact, we use sediment streams, shown in Figure 1, as an indicator of benthic feeding behavior when analyzing drone footage (Torres et al. 2018).
Locations of benthic feeding can be identified without directly observing a gray whale actively feeding because of the excavated pits that result from benthic feeding (Nerini 1984). These pits can be detected using side-scan sonar that is commonly used to map the seafloor. Oliver and Slattery (1985) found that the pits typically are from 2-20 m2. In some of the imagery, consecutive neighboring pits are visible, likely created by one whale in series during a feeding event. Figure 2 shows different arrangements of pits.
Aside from how fascinating the behavior is, benthic feeding is also interesting because it has a large impact on the environment. Coming from a background of studying baleen whales that primarily feed on krill, I had not really considered the potential impacts of whale foraging other than removing prey from the environment. However, when gray whales feed, they excavate large areas of the benthic substrate that disturb and impact the habitat.
The impacts of benthic feeding
Weitkamp et al. (1992) conducted a study on gray whale benthic foraging on ghost shrimp in Puget Sound, WA, USA. This study, conducted over two years, focused on measuring the impact of benthic foraging by its effect on prey abundance. They found that the standing stock of ghost shrimp within a recently excavated pit was two to five times less than that outside the pit, and that 3100 to 5700 grams of shrimp can be removed per pit. From aerial surveys they estimated that within one season feeding gray whales created between 2700 and 3200 pits. Using these values, they calculated that 55 to 79% of the standing stock of ghost shrimp was removed each season by foraging gray whales. Interestingly, they found that the shrimp biomass within an excavated pit recovered within about two months.
Oliver and Slattery (1985) also
found a recovery period of about 2 months per pit in their study on the effect
of gray whale benthic feeding on the prey community in the Bering Sea. They
sampled prey within and outside feeding excavations, both actual whale pits and
man-made, to test the response of the benthic community to the disturbance of a
feeding event. They found that after the initial feeding disturbance, the
excavated area was rapidly colonized by scavenging lysianassid amphipods, which
are small (10 mm) crustaceans that typically eat dead organic material. These
amphipods rushed in and attacked the organisms that were injured or dislodged
by the whale feeding event, typically small crustaceans and polychaete worms.
Within hours of the whale feeding event, these amphipods had dispersed and a
different genre of scavenging lysianassid amphipods slowly invaded the
excavated pit further and stayed much longer. After a few days or weeks these
pits collected and trapped organic debris that attracted more colonists.
Indeed, they found that the number of colonists remained elevated within the
excavated areas for over two months.
Notably, these results on how the
disturbance of gray whale benthic feeding changes sediment composition support
the idea that this foraging behavior maintains the sand substrate and therefore
helps to maintain balanced levels of benthic dwelling amphipods, their primary
source of prey in this study area (Johnson and Nelson, 1984). Gray whales scour
the sea floor when they feed and this process leads to the resuspension of lots
of sediments and nutrients that would otherwise remain on the seafloor.
Therefore, while this feeding may seem like a violent disturbance, it may in
fact play a large role in benthic productivity (Johnson and Nelson, 1984;
Oliver and Slattery, 1985).
These ecosystem impacts of gray
whale benthic feeding I have described above demonstrate the various stages of
invaders after a feeding disturbance, and the process of succession. Succession
is the ecological process of how a community structure builds and grows.
Primary succession is when the structure grows from truly nothing and secondary
succession occurs after a disturbance, such as a fire. In secondary succession,
there are typically pioneer species that first appear and then give way to
other species and a more complex community eventually emerges. Succession is
well documented in many terrestrial studies after disturbance events, and the
processes of secondary succession is very important to community ecology and
Since gray whale benthic foraging
does not impact an entire habitat all at once, the process is not perfectly
comparable to secondary succession in terrestrial systems. Yet, when thinking
about the smaller scale, another example of succession in the marine environment
takes place at a whale fall. When a whale dies and sinks to the ocean floor, a
small ecosystem emerges. Different organisms arrive at different stages to
scavenge different parts of the carcass and a food web is created around it.
me the impacts of gray whale benthic feeding are akin to both terrestrial disturbance
events and whale falls. The excavation serves as a disturbance, and through secondary
succession the habitat is refreshed via stages of different species colonization
until the system eventually returns to the pre-disturbance levels. However,
like a whale fall the feeding event leaves behind injured or displaced
organisms that scavengers consume; in fact seabirds are known to take advantage
of benthic invertebrates that are brought to the surface by a gray whale feeding
event (Harrison, 1979).
So much of our research is focused
on questions about how the changing environment impacts our study species and
not the other way around. This venture into the literature has provided me with
an important reminder to think about flipping the question. I have enjoyed
starting 2020 with a reminder of how cool gray whales are, and that while a
disturbance can initially be thought of as negative, it may actually bring
about important, and positive, change.
Nerini, Mary. 1984. “A Review of Gray Whale Feeding
Ecology.” In The Gray Whale: Eschrichtius Robustus, 423–50. Elsevier
Oliver, J. S., and P. N. Slattery. 1985. “Destruction and
Opportunity on the Sea Floor: Effects of Gray Whale Feeding.” Ecology 66
(6): 1965–75. https://doi.org/10.2307/2937392.
Torres, Leigh G., Sharon L. Nieukirk, Leila Lemos, and Todd
E. Chandler. 2018. “Drone up! Quantifying Whale Behavior from a New Perspective
Improves Observational Capacity.” Frontiers in Marine Science 5 (SEP).
Weitkamp, Laurie A, Robert C Wissmar, Charles A Simenstad,
Kurt L Fresh, and Jay G Odell. 1992. “Gray Whale Foraging on Ghost Shrimp
(Callianassa Californiensis) in Littoral Sand Flats of Puget Sound, USA.” Canadian
Journal of Zoology 70 (11): 2275–80. https://doi.org/10.1139/z92-304.
Johnson, Kirk R., and C. Hans Nelson. 1984. “Side-Scan Sonar
Assessment of Gray Whale Feeding in the Bering Sea.” Science 225 (4667):
Harrison, Craig S. 1979. “The Association of Marine Birds
and Feeding Gray Whales.” The Condor 81 (1): 93.
In the GEMM Lab, our research focuses largely on the ecology of marine top predators. Inherent in our work are often assumptions that our study species—wide-ranging predators including whales, dolphins, otters, or seabirds—will distribute themselves relative to their prey. In order to make a living in the highly patchy and dynamic marine environment, predators must find ways to predictably locate and exploit prey resources.
But what about the prey? How do the prey structure themselves relative to their predators? This question is explored in depth in a paper titled “The Landscape of Fear: Ecological Implications of Being Afraid” (Laundre et al. 2010), which we discussed in our most recent lab meeting. When wolves were re-introduced in Yellowstone, the elk increased their vigilance and altered their grazing patterns. As a result, the plant community was altered to reflect this “landscape of fear” that the elk move through, where their distribution not only reflected opportunities for the elk to eat but also the risk of being eaten.
Translating the landscape of fear concept to the marine environment is tricky, but a fascinating exercise in ecological theory. We grappled with drawing parallels between the example system of wolves, elk, and vegetation and baleen whales, zooplankton, and phytoplankton. Relative to grazing mammals like elk, the cognitive abilities of zooplankton like krill, copepods, and mysid might pale in comparison. How could we possibly measure “fear” or “vigilance” in zooplankton? The swarming behavior of mysid and krill into dense patches is a defense mechanism—the strategy they have evolved to lessen the likelihood that any one of them will be eaten by a predator. I would posit that the diel vertical migration (DVM) of zooplankton is a manifestation of fear, at least on some level. DVM occurs over the course of each day, with plankton in pelagic ecosystems migrating vertically in the water column to avoid predators by hiding at depth during the daylight hours, and then swimming upward to feed on phytoplankton under the cover of darkness. I won’t speculate any further on the intelligence of zooplankton, but the need to survive predation has driven them to evolve this effective evolutionary strategy of hiding in the ocean’s twilight zone, swimming upward to feed only after dark so that they’re less likely to linger in spaces occupied by predators.
Laundre et al. (2010) present a visual representation of the landscape of fear (Fig. 1, reproduced below), where as an animal moves through space (represented as distance in meters or kilometers, for example), they also move through varying levels of predation risk. Environmental gradients (temperature, for example) tend to be much more stable across space in terrestrial ecosystems such as in the Yellowstone example from the paper. I wonder whether the same concept and visual depiction of a landscape of fear could be translated as risk across various environmental gradients, rather than geographic distances? In this proposed illustration, a landscape of fear would vary based on gradients of environmental conditions rather than geographic space. Such a shift in spatial reference —from geographic to environmental space—might make the model more applicable in the dynamic ocean ecosystems that we study.
What about cases when the predators we study become prey? One example we discussed was gray whales migrating from breeding lagoons in Mexico to feeding grounds in the Bering Sea. Mother-calf pairs hug the coastline tightly, by no means taking the most direct route between locations and adding considerable travel distance to their migration. The leading hypothesis is that mother gray whales take the coastal route to minimize the risk that their calves will fall prey to killer whale attacks. Are there other cases where the predators we study operate in a seascape of fear that we do not yet understand? Likely so, and the predators’ own seascape of fear may account for cases when we cannot explain predator distribution simply by their prey and their environment. To take this a step further, it might be beneficial not only to think of predation risk as only the potential to be eaten, but expand our definition to include human disturbance. While humans may not directly prey on marine predators, the disturbance from human activity in the ocean likely creates a layer of fear which animals must navigate, even in the absence of actual predation.
Our lively lab meeting discussion prompted me to look into
how the landscape of fear model has been applied to the highly dynamic and
intricate marine environment. In a study examining predator-prey dynamics of
three species of marine mammals—bottlenose dolphins, harbor seals, and
dugongs—Wirsing et al. (2007) found that in all three cases, the study species
spent less time in more desirable prey patches or decreased riskier behavior in
the presence of predators. Most studies in marine ecology are observational, as
we rarely have the opportunity to manipulate our study system for experimental
design and hypothesis testing. However, a study of coral reefs in the Florida
Keys conducted by Catano et al. (2015) used fabricated predators—decoys of
black grouper, a predatory fish—to investigate the influence of fear of
predation on the reef system. What they found was that herbivorous fish
consumed significantly less and fed at a much faster rate in the presence of this
decoy predator. The grouper, even in decoy form, created a “reefscape of fear”,
altering patterns in herbivory with potential ramifications for the entire ecosystem.
My takeaway from our discussion and my musings in this
week’s blog post is that predator and prey distribution and behavior is highly
interconnected. While predators distribute themselves to maximize their ability
to find a meal, their prey respond accordingly by balancing finding a meal of
their own with minimizing the risk that they will be eaten. Ecology is the
study of an ecosystem, which means the questions we ask are complicated and
hierarchical, and must be considered from multiple angles, accounting for
biological, environmental, and behavioral elements to name a few. These
challenges of studying ecosystems are simultaneously what make ecology
fascinating, and exciting.
Laundré, J. W., Hernández, L., & Ripple, W. J. (2010).
The landscape of fear: ecological implications of being afraid. Open
Ecology Journal, 3, 1-7.
Catano, L. B., Rojas, M. C., Malossi, R. J., Peters, J. R.,
Heithaus, M. R., Fourqurean, J. W., & Burkepile, D. E. (2016). Reefscapes
of fear: predation risk and reef hetero‐geneity interact to shape herbivore
foraging behaviour. Journal of Animal Ecology, 85(1),
Wirsing, A. J., Heithaus, M. R., Frid, A., & Dill, L. M.
(2008). Seascapes of fear: evaluating sublethal predator effects experienced
and generated by marine mammals. Marine Mammal Science, 24(1),
By: Alexa Kownacki, Ph.D. Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab
Data analysis is often about parsing down data into manageable subsets. My project, which spans 34 years and six study sites along the California coast, requires significant data wrangling before full analysis. As part of a data analysis trial, I first refined my dataset to only the San Diego survey location. I chose this dataset for its standardization and large sample size; the bulk of my sightings, over 4,000 of the 6,136, are from the San Diego survey site where the transect methods were highly standardized. In the next step, I selected explanatory variable datasets that covered the sighting data at similar spatial and temporal resolutions. This small endeavor in analyzing my data was the first big leap into understanding what questions are feasible in terms of variable selection and analysis methods. I developed four major hypotheses for this San Diego site.
H1: I predict that bottlenose dolphin sightings along the San Diego transect throughout the years 1981-2015 exhibit clustered distribution patterns as a result of the patchy distributions of both the species’ preferred habitats, as well as the social nature of bottlenose dolphins.
H2: I predict there would be higher densities of bottlenose dolphin at higher latitudes spanning 1981-2015 due to prey distributions shifting northward and less human activities in the northerly sections of the transect.
H3: I predict that during warm (positive) El Niño Southern Oscillation (ENSO) months, the dolphin sightings in San Diego would be distributed more northerly, predominantly with prey aggregations historically shifting northward into cooler waters, due to (secondarily) increasing sea surface temperatures.
H4: I predict that along the San Diego coastline, bottlenose dolphin sightings are clustered within two kilometers of the six major lagoons, with no specific preference for any lagoon, because the murky, nutrient-rich waters in the estuarine environments are ideal for prey protection and known for their higher densities of schooling fishes.
The common bottlenose dolphin (Tursiops truncatus) sighting data spans 1981-2015 with a few gap years. Sightings cover all months, but not in all years sampled. The same transect in San Diego was surveyed in a small, rigid-hulled inflatable boat with approximately a two-kilometer observation area (one kilometer surveyed 90 degrees to starboard and port of the bow).
I wanted to see if there were changes in dolphin distribution by latitude and, if so, whether those changes had a relationship to ENSO cycles and/or distances to lagoons. For ENSO data, I used the NOAA database that provides positive, neutral, and negative indices (1, 0, and -1, respectively) by each month of each year. I matched these ENSO data to my month-date information of dolphin sighting data. Distance from each lagoon was calculated for each sighting.
H1:True, dolphins are clustered and do not have a uniform distribution across this area. Spatial analysis indicated a less than a 1% likelihood that this clustered pattern could be the result of random chance (Fig. 1, z-score = -127.16, p-value < 0.0001). It is well-known that schooling fishes have a patchy distribution, which could influence the clustered distribution of their dolphin predators. In addition, bottlenose dolphins are highly social and although pods change in composition of individuals, the dolphins do usually transit, feed, and socialize in small groups.
H2:False, dolphins do not occur at higher densities in the higher latitudes of the San Diego study site. The sightings are more clumped towards the lower latitudes overall (p < 2e-16), possibly due to habitat preference. The sightings are closer to beaches with higher human densities and human-related activities near Mission Bay, CA. It should be noted, that just north of the San Diego transect is the Camp Pendleton Marine Base, which conducts frequent military exercises and could deter animals.
H3: False, during warm (positive) El Niño Southern Oscillation (ENSO) months, the dolphin sightings in San Diego were more southerly. In colder (negative) ENSO months, the dolphins were more northerly. The differences between sighting latitude and ENSO index was significant (p<0.005). Post-hoc analysis indicates that the north-south distribution of dolphin sightings was different during each ENSO state.
H4:True, dolphins are clustered around particular lagoons. Figure 5 illustrates how dolphin sightings nearest to Lagoon 6 (the San Dieguito Lagoon) are always within 0.03 decimal degrees. Because of how these data are formatted, decimal degrees is the easiest way to measure change in distance (in this case, the difference in latitude). In comparison, dolphins at Lagoon 5 (Los Penasquitos Lagoon) are distributed across distances, with the most sightings further from the lagoon.
I found a significant difference between distance to nearest lagoon in different ENSO index categories (p < 2.55e-9): there is a significant difference in distance to nearest lagoon between neutral and negative values and positive and neutral years. Therefore, I hypothesize that in neutral ENSO months compared to positive and negative ENSO months, prey distributions are changing. This is one possible hypothesis for the significant difference in lagoon preference based on the monthly ENSO index. Using a violin plot (Fig. 6), it appears that Lagoon 5, Los Penasquitos Lagoon, has the widest variation of sighting distances in all ENSO index conditions. In neutral years, Lagoon 0, the Buena Vista Lagoon has multiple sightings, when in positive and negative years it had either no sightings or a single sighting. The Buena Vista Lagoon is the most northerly lagoon, which may indicate that in neutral ENSO months, dolphin pods are more northerly in their distribution.
Takeaways to science and management:
Bottlenose dolphins have a clustered distribution which seems to be related to ENSO monthly indices, and likely, their social structures. From these data, neutral ENSO months appear to have something different happening compared to positive and negative months, that is impacting the sighting distributions of bottlenose dolphins off the San Diego coastline. More research needs to be conducted to determine what is different about neutral months and how this may impact this dolphin population. On a finer scale, the six lagoons in San Diego appear to have a spatial relationship with dolphin sightings. These lagoons may provide critical habitat for bottlenose dolphins and/or for their preferred prey either by protecting the animals or by providing nutrients. Different lagoons may have different spans of impact, that is, some lagoons may have wider outflows that create larger nutrient plumes.
Other than the Marine Mammal Protection Act and small protected zones, there are no safeguards in place for these dolphins, whose population hovers around 500 individuals. Therefore, specific coastal areas surrounding lagoons that are more vulnerable to habitat loss, habitat degradation, and/or are more frequented by dolphins, may want greater protection added at a local, state, or federal level. For example, the Batiquitos and San Dieguito Lagoons already contain some Marine Conservation Areas with No-Take Zones within their reach. The city of San Diego and the state of California need better ways to assess the coastlines in their jurisdictions and how protecting the marine, estuarine, and terrestrial environments near and encompassing the coastlines impacts the greater ecosystem.
This dive into my data was an excellent lesson in spatial scaling with regards to parsing down my data to a single study site and in matching my existing data sets to other data that could help answer my hypotheses. Originally, I underestimated the robustness of my data. At first, I hesitated when considering reducing the dolphin sighting data to only include San Diego because I was concerned that I would not be able to do the statistical analyses. However, these concerns were unfounded. My results are strongly significant and provide great insight into my questions about my data. Now, I can further apply these preliminary results and explore both finer and broader scale resolutions, such as using the more precise ENSO index values and finding ways to compare offshore bottlenose dolphin sighting distributions.
By Dominique Kone, Masters Student in Marine Resource Management
Species reintroductions are a management strategy to augment the reestablishment or recovery of a locally-extinct or extirpated species into once native habitat. The potential for reestablishment success often depends on the species’ ecological characteristics, habitat requirements, and relationship and effects to other species in the environment. While the science behind species reintroductions is continuously evolving and improving, reintroductions are still inherently risky and uncertain in nature. Therefore, every effort should be made to fully assess ecological factors before a reintroduction takes place. As Oregon considers a potential sea otter reintroduction, understanding these ecological factors is an important piece of my own graduate research.
Sea otters are oftentimes referred to as keystone species because they can have wide-reaching effects on the community structure and function of nearshore marine environments. Furthermore, relative to other marine mammals or top predators, several papers have documented these effects – partially due to the ease in observing their foraging and social behaviors, which typically take place close to shore. In many of these studies, a classic paradigm repeatedly appears: when sea otters are present, prey densities (e.g., sea urchins) are significantly reduced, while macroalgae (e.g., kelp, seagrass) densities are high.
While this paradigm is widely-accepted amongst researchers, a few key studies have also demonstrated that the effects of sea otters may be more variable than we once thought. The paradigm does not necessarily hold true everywhere sea otters exist, or at least not to the same degree. For example, after observing benthic communities along islands with varying sea otter densities in the Aleutian archipelago, Alaska, researchers found that islands with abundant otter populations consistently supported low sea urchin densities and high, yet variable, kelp densities. In contrast, islands without otters consistently had low kelp densities and high, yet variable, urchin densities. This study demonstrates that while the classic paradigm generally held true, the degree to which the ecosystem belonged to one of two dominant states (sea otters, low urchins, and high kelp or no sea otters, high urchins, and low kelp) was less obvious.
This example demonstrates the danger in applying this one-size-fits-all paradigm to sea otter effects. Hence, we want to achieve a better understanding of potential sea otter effects so that managers may anticipate how Oregon’s nearshore environments may be affected if sea otters were to be reintroduced. Yet, how can we accurately anticipate these effects given these potential variations and deviations from the paradigm? Interestingly, if we look to other fields outside ecology, we find a possible solution and tool for tackling these uncertainties: a systematic review of available literature.
For decades, medical researchers have been conducting systematic reviews to assess the efficacy of treatments and drugs by combining several studies to find common findings. These findings can then be used to determine any potential variation between studies (i.e. instances where the results may conflict or differ from one another) and even test the influence and importance of key factors that may be driving that variation. While systematic reviews are quite popular within the medical research field, they have not been applied regularly in ecology, but recognition of their application to ecological questions is growing. In our case of achieving a better understanding of the drivers of ecological impacts of sea otter, a systematic literature review is an ideal tool to assess variable effects. This review will be the focus of my second thesis chapter.
In conducting my review, there will be three distinct phases: (1) review design and study collection, (2) meta-analysis, and (3) factor testing. In the first phase (review design and study collection), I will search the existing literature to collect studies that explicitly compare the availability of key ecosystem components (i.e. prey species, non-prey species, and macroalgae species) when sea otters are absent and present in the environment. By only including studies that make this comparison, I will define effects as the proportional change in each species’ or organism group’s availability (e.g. abundance, biomass, density, etc.) with and without sea otters. In determining these effects, it’s important to recognize that sea otters alter ecosystems via both direct and indirect pathways. Direct effects can be thought of as any change to prey availability via sea otter predation directly, while indirect effects can be thought of an any alteration to the broader ecosystem (i.e. non-prey species, macroalgae, habitat features) as an indirect result from sea otter predation on prey species. I will record both types of effects.
In phase two, I will use meta-analytical procedures (i.e. statistical analyses specific to systematic reviews) to calculate one standardized metric to represent sea otter effects. These effects will be calculated and averaged across all collected studies. As previously discussed, there may be key factors – such as sea otter density – that influence these effects. Therefore, in phase three (factor testing), effects will also be calculated separately for each a priori factor to test their influence on the effects. Such factors may include habitat type (i.e. hard or soft sediment), prey species (i.e. sea urchins, crabs, clams, etc.), otter density, depth, or time after otter recolonization.
In statistical terms, the goal of testing factors is to see if the variation between studies is impacted by calculating sea otter effects separately for each factor versus across all studies. In other words, if we find high variation in effects between studies, there may be important factors driving that variation. Therefore, in systematic reviews, we recalculate effects separately for each factor to try to explain that variation. If, however, after testing these factors, variation remains high, there may be other factors that we didn’t test that could be driving that remaining variation. Yet, without a priori knowledge on what those factors could be, such variation should be reported as a major source of uncertainty.
Predicting or anticipating the effects of reintroduced species is no easy feat. In instances where the ecological role of a species is well known – and there is adequate data – researchers can develop and use ecosystem models to predict with some certainty what these effects may be. Yet, in other cases where the species’ role is less studied, has less data, or is more variable, researchers must look to other tools – such as systematic reviews – to gain a better understanding of these potential effects. In this case, a systematic review on sea otter effects may prove particularly useful in helping managers understand what types of ecological effects of sea otters in Oregon are most likely, what the important factors are, and, after such review, what we still don’t know about these effects.
 Seddon, P. J., Armstrong, D. P., and R. F. Maloney. 2007. Developing the science of reintroduction biology. Conservation Biology. 21(2): 303-312.
 Estes, J. A., Tinker, M. T., and J. L. Bodkin. 2009. Using ecological function to develop recovery criteria for depleted species: sea otters and kelp forests in the Aleutian Archipelago. Conservation Biology. 24(3): 852-860.
 Sutton, A. J., and J. P. T. Higgins. 2008. Recent developments in meta-analysis. Statistics in Medicine. 27: 625-650.
 Arnqvist, G., and D. Wooster. 1995. Meta-analysis: synthesizing research findings in ecology and evolution. TREE. 10(6): 236-240.
 Vetter, D., Rucker, G., and I. Storch. 2013. Meta-analysis: a need for well-defined usage in ecology and conservation biology. Ecosphere. 4(6): 1-13.
By Dominique Kone, Masters Student in Marine Resource Management
A couple of months ago, I wrote a blog introducing our new project, and my thesis, on the potential to reintroduce sea otters to the Oregon coast. In that blog, I expressed that in order to develop a successful reintroduction plan, scientists and managers need to have a sound understanding of sea otter ecology and the current state of Oregon’s coastal ecosystems. As a graduate student conducting a research-based thesis in a management program, I’m constantly fretting over the applicability of my research to inform decision-making processes. However, in the course of conducting my research, I sometimes forget just how COOL sea otters are. Therefore, in this blog, I wanted to take the opportunity to nerd out and provide you with my top five favorite facts about these otterly adorable creatures.
Without further ado, here are my top five favorite facts about sea otters:
Sea otters eat a lot. Previous studies show that an individual sea otter eats up to 30% of its own body weight in food each day. With such high caloric demands, sea otters spend a great deal of their time foraging the seafloor for a variety of prey species, and have been shown to decrease prey densities in their local habitat significantly. Sea otters are famously known for their taste for sea urchins. Yet, these voracious predators also consume clams, sea stars, crabs, and a variety of other small invertebrate species.
Individuals are specialists, but can change their diet. Sea otters typically show individual foraging specialization – which means an individual predominantly eats a select few species of prey. However, this doesn’t mean an otter can’t switch or consume other types of prey as needed. In fact, while individuals tend to be specialists, on a population or species level, sea otters are actually generalist predators. Past studies that looked at the foraging habits of expanding sea otter populations show that as populations expand into unoccupied territory, they typically eat a limited number of prey. But as populations grow and become more established, the otters will start to diversify their diet, suggesting intra-specific competition.
Sea otters exert a strong top-down force. Top-down forcing is one of the most important concepts we must acknowledge when discussing sea otter ecology. With top-down forcing, consumers at the top of the food chain depress the trophic level on which they feed, and this feeding indirectly increases the abundance of the next lower trophic level, resulting in a cascading effect. The archetype example of this phenomenon is the relationship between sea otters, sea urchins, and kelp forests. This relationship goes as follows: sea otters consume sea urchin, and sea urchins graze on kelp. Therefore, sea otters reduce sea urchin densities by direct predation, thereby mediating grazing pressure on kelp. This indirect effect allows kelp to grow more abundantly, which is why we often see relatively productive kelp forests when sea otters are present. This top-down forcing also has important implications for the whole ecosystem, as I’ll explain in my next fact.
Sea otters help restore ecosystems, and associated ecosystem services. In kelp habitat where sea otters have been removed, we often see high densities of sea urchins and low biomasses of kelp. In this case, sea urchins have no natural predators to keep their populations in check and therefore completely decimate kelp forests. However, what we’ve learned is that when sea otters “reclaim” previously occupied habitats or expand into unoccupied territory, they can have remarkable restorative effects because their predation on sea urchins allows for the regrowth of kelp forest. Additionally, with the restoration of key ecosystems like kelp forests, we can see a variety of other indirect benefits – such as increased biodiversity, refuge for fish nurseries and commercially-important species, and carbon sequestration. The structure of nearshore ecosystems and communities change drastically with the addition or removal of sea otters, which is why they’re often referred to as keystone species.
Sea otters are most often associated with coastal kelp forests, but they can also exist in other types of habitats and ecosystems. In addition to kelp dominated ecosystems, sea otters are known to use estuaries and bays, seagrass beds, and swim over a range of bottom substrates. As evidenced by previous studies, sea otters exert similar top-down forces in non-kelp ecosystems, as they do within kelp forests. One study found that sea otters also had restorative effects on seagrass beds within estuaries, where they consumed different types of prey (i.e., crabs instead of urchins), demonstrating that sea otters play a significant keystone role in seagrass habitats as well . Findings such as these are vitally important to understanding (1) where sea otters are capable of living relative to habitat characteristics, and (2) how recovering or expanding sea otter populations may impact ecosystems and habitats in which they don’t currently exist, such as the Oregon coast.
Well, there you have it – my top five favorite facts about sea otters. This list is by no means exhaustive of all there is to know about sea otter ecology, and isn’t enough information to develop an informative reintroduction plan. However, a successful reintroduction plan will rely heavily on these underlying ecological characteristics of sea otters, in addition to the current state of Oregon’s nearshore ecosystems. As someone who constantly focuses on the relationship between scientific research and management and conservation, it’s nice every now and then to take a step back and just simply appreciate sea otters for being, well, sea otters.
 Costa, D. P. 1978. The ecological energetics, water, and electrolyte balance of the California sea otter (Enhydra lutris). Ph.D. dissertation, University of California, Santa Cruz.
 Reidman, M. L. and J. A. Estes. 1990. The sea otter (Enhydra lutris): behavior, ecology, and natural history. United States Department of the Interior, Fish and Wildlife Service, Biological Report. 90: 1-126.
 Laidre, K.L. and R. J. Jameson. 2006. Foraging patterns and prey selection in an increasing and expanding sea otter population. Journal of Mammology. 87(4): 799-807.
 Estes, J. A., Jameson, R.J., and B. R. Rhode. 1982. Activity and prey election in the sea otter: influence of population status on community structure. The American Naturalist. 120(2): 242-258.
 Tinker, M. T., Costa, D. P., Estes, J. A., and N. Wieringa. 2007. Individual dietary specialization and dive behavior in the California sea otter: using archival time-depth data to detect alternative foraging strategies. Deep-Sea Research Part II. (54):330-342.
 Newsome et al. 2009. Using stable isotopes to investigate individual diet specialization in California sea otters (Enhydra lutris nereis). Ecology. 90(4): 961-974.
 Ostfeld, R. S. 1982. Foraging strategies and prey switching in the California sea otter. Oecologia. 53(2): 170-178.
 Paine, R. T. 1980. Food webs: linkage, interaction strength and community infrastructure. The Journal of Animal Ecology. 49(3): 666-685.
 Estes, J. A. and J.F. Palmisano. 1974. Sea otters: their role in structuring nearshore communities. Science. 185(4156): 1058-1060.
 Estes, J. A., and D. O. Duggins. 1995. Sea otters and kelp forests in Alaska: generality and variation in a community ecological paradigm. Ecological Monographs. 65(1): 75-100.
 Wilmers, C. C., Estes, J. A., Edwards, M., Laidre, K. L., and B. Konar. 2012. Do trophic cascades affect the storage and flux of atmospheric carbon? An analysis of sea otters and kelp forests. Frontiers in Ecology and the Environment. 10(8): 409-415.
 Hughes et al. 2014. Recovery of a top predator mediate negative eutrophic effects on seagrass. Proceedings of the National Academy of Sciences. 110(38): 15313-15318.
 Lee, L.C., Watson, J. C., Trebilco, R., and A. K. Salomon. Indirect effects and prey behavior mediate interactions between an endangered prey and recovering predator. Ecosphere. 7(12).
 Laidre, K. L., Jameson, R. J., Gurarie, E., Jeffries, S. J., and H. Allen. 2009. Spatial habitat use patterns of sea otters in coastal Washington. Journal of Mammalogy. 90(4): 906-917.
 Lafferty, K. D., and M. T. Tinker. 2014. Sea otters are recolonizing southern California in fits and starts. Ecosphere. 5(5).