Are there picky eaters in the PCFG?



















Connecting Research Questions
















Milling around in definitions


































The complex relationship between behavior and body condition
















What are the ecological impacts of gray whale benthic feeding?


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 feed?

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).

Figure 1. Screenshot of drone video showing sediment streaming from mouth of a whale after benthic feeding. Video taken under NOAA/NMFS permit #21678

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.

Figure 2. Different arrangements of pits created by feeding whales (Nerini 1984).

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 resilience.

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.

To 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 Inc.

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.


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.

Johnson, Kirk R., and C. Hans Nelson. 1984. “Side-Scan Sonar Assessment of Gray Whale Feeding in the Bering Sea.” Science 225 (4667): 1150–52.

Harrison, Craig S. 1979. “The Association of Marine Birds and Feeding Gray Whales.” The Condor 81 (1): 93.

Classifying cetacean behavior



How do we collect data on behavior?




Categorizing Behaviors







































Midway Atoll: the next two weeks at the largest albatross colony in the world (two years later)


This February I had the opportunity to spend two weeks at Midway Atoll National Wildlife Refuge in the Papahānaumokuākea Marine National Monument. I was there to GPS track black-footed and Laysan albatross during their short chick-brooding foraging trips. Two weeks is just enough time since the albatross are taking short trips (3-5 days) to feed their rapidly growing chicks.

My first visit to Midway (2016 blog post) occurred right as the black-footed albatross chicks were hatching (quickly followed by the Laysan albatross chicks). This time, we arrived almost exactly when I had left off. The oldest chicks were just about two weeks old. This shift in phenology meant that, though subtle, each day offered new insights for me as I watched chicks transform into large aware and semi-mobile birds. By the time we left, unattended chicks were rapidly multiplying as the adults shifted to the chick-rearing stage. During chick rearing, both parents leave the chick unattended and take longer foraging trips.

Our research goal was to collect tracking data from both species that can be used to address a couple of research questions. First of all, winds can aid, or hinder albatross foraging and flight efficiency (particularly during the short brooding trips). In the North Pacific, the strength and direction of the winds are influenced by the ENSO (El Niño Southern Oscillation) cycles. The day after we left Midway, NOAA issued an El Niño advisory indicating weak El Nino conditions. We know from previous work at Tern Island (farther east and farther south at 23.87 N, -166.28 W) that El Niño improves foraging for Laysan albatrosses during chick brooding, while during La Niña reproductive success is lower (Thorne et al., 2016). However, since Midway is farther north, and farther west the scenario might be different there. Multiple years of GPS tracking data are needed to address this question and we hope to return to collect more data next year (especially if  La Niña follows the El Niño as is often the case).

We will also overlap the tracking data with fishing boat locations from the Global Fishing Watch database to assess the potential for birds from Midway to interact with high seas fisheries during this time of year (project description, associated blog post). Finally, many of the tags we deployed incorporated a barometric pressure sensor and the data can be used to estimate flight heights relative to environmental conditions such as wind strength. This type of data is key to assessing the impact of offshore wind energy (Kelsey et al., 2018).

How to track an albatross

To track an albatross we use small GPS tags that we tape to the back feathers. After the bird returns from a foraging trip, we remove the tape from the feathers and take the datalogger off. Then we recharge the battery and download the data!

This research is a collaboration between Lesley Thorne (Stony Brook University), Scott Shaffer (San Jose State University), myself (Oregon State University), and Melinda Conners (Washington State University). The field effort was generously supported by the Laurie Landeau Foundation via the Minghua Zhang Early Career Faculty Innovation Fund at Stoney Brook University to Lesley Thorne.

My previous visit to Midway occurred just after house mice were discovered attacking incubating adult albatrosses. Since then, a lot of thought and effort had gone into developing a plan to eradicate mice from Midway. You can find out more via Island Conservation’s Midway blogs and the USFWS.


Kelsey, E. C., Felis, J. J., Czapanskiy, M., Pereksta, D. M., & Adams, J. (2018). Collision and displacement vulnerability to offshore wind energy infrastructure among marine birds of the Pacific Outer Continental Shelf. Journal of Environmental Management, 227, 229–247.

Thorne, L. H., Conners, M. G., Hazen, E. L., Bograd, S. J., Antolos, M., Costa, D. P., & Shaffer, S. A. (2016). Effects of El Niño-driven changes in wind patterns on North Pacific albatrosses. Journal of the Royal Society Interface, 13(119), 20160196.

Who Am I? Exploring the theory of individualisation among marine mammals


“Just be yourself!” is a phrase that everyone has probably heard at least once in their lives. The idea of being an individual who is distinctly different from other individuals is a concept that is focal to the society we live in today. While historically it may have been frowned upon to be the “black sheep in the crowd”, nowadays that seems to be the goal.

Source: Go Comics.

This quest for uniqueness has resulted in different styles of fashion, speech, profession, interest in art, music, literature, automobile types – the list is endless. The American Psychological Association defines personality as the “individual differences in characteristic patterns of thinking, feeling and behaving”1. So, all of the choices we make on a daily basis shape our behaviour, and our behaviour in turn shapes our personality.

Since personality is something that is so engrained within human society, it isn’t surprising that ecologists have explored this concept among non-humans. Decades of research have resulted in an abundance of literature detailing personality in many different taxa and species, ranging from chimpanzees to mice to ants2. Naturally, the definition of personality for animals differs from that for humans since the assessment of animal thoughts and feelings is still somewhat of a locked box to us. Nevertheless, the behavioural aspect of the two definitions remains consistent whereby animal personality is broadly defined as “consistent variation in behavioural traits between individuals”3.

Although I am an early career marine mammal ecologist finding my footing in this rapidly expanding field, I have a keen interest in teasing apart possible cases of individual specialisation within marine mammal populations. So, before getting straight into the nitty gritty of individual specialisation, it is important for me to take a small step back and consider the concept of specialisation as applied to small subgroups or populations of marine mammals.

Specialisations are mostly related to foraging or feeding behaviour whereby a subgroup of individuals will develop a novel method to locate and capture prey. These behaviours have been reported for several marine mammal species, and are strongly coupled to intra and inter-specific competition with other predators for prey and habitat characteristics. Furthermore, it is posited that factors such as resource benefits (e.g. energy content of prey), prey escape rates, and handling times can be minimised if specialisation for a particular prey type or habitat occurs4.

In Florida Bay, Torres & Readdocumented two distinct foraging strategies employed by two bottlenose dolphin ecotypes. One dolphin ecotype was found to forage using deep diving with erratic surfacings, whereas the second ecotype chose to forage through mud ring feeding and were mostly seen in shallow habitats. The latter ecotype is in fact so adapted to shallow depths that dolphins were typically observed foraging in waters <2 m deep. In this example, the foraging tactics of the two ecotypes are strongly driven by habitat conditions, specifically depth. The video below is aerial footage of bottlenose dolphins performing mud ring feeding.

Such group specialisations have been identified not only in several other bottlenose dolphin populations around the world6,7, but also in other cetacean species, including killer whales (distinct differences in target prey between transients and residents8), Guiana dolphins (mud-plume feeding9), humpback dolphins (strand feeding10), and several others. Noticeable here is that these records concern Odontocete species, which is not surprising since these toothed whales are vastly different to baleen whales in that they often live in structured groups with bonds between individuals sometimes lasting for decades11. Long-term relationships are conducive to developing specialised group hunting strategies as individuals will spend considerable time with one another and the success of obtaining prey depends on the cooperation and coordination of the group.

For baleen whales and other marine mammals, such as pinnipeds, where life history and social organisation is more geared toward a solitary life, examples of group specialisations are relatively rare (with the exception of the well-documented bubble-net feeding exhibited by humpback whales12). While group specialisation may not be as prevalent in Mysticetes, the same problems of inter and intra-specific competition persists among these more solitary species too, which would suggest that individuals should develop their own unique foraging tactics and preferences. Evidence for individualisation is hard to obtain since it requires repeated observations of the same individuals over time with good knowledge of the prey type being consumed and/or the habitat being used to forage in.

Nevertheless, examples do exist. Perhaps the most well-documented case of individualisation within a population for marine mammals is of the sea otter. Estes et al. (2003) describe 10 female sea otters in Monterey Bay that had high inter-individual variation in diet, which they investigated over a scale of 8 years13. Most females specialised on 1-4 types of prey, with marked differences between the diets chosen by each female, despite habitat overlap. This individualisation of diet was not attributable to variation in prey availability; hence, authors concluded that this extreme specialisation occurred to reduce intra-population competition for prey.

Ecologists have historically (and probably still to this day) disagreed on whether individualisation actually matters in the grand scheme of things. There are generally three schools of thought on the matter: (1) individual specialisation is rare and/or weakly influences population dynamics and so is not very important; (2) while individual specialisation does occur and may in fact be commonplace, it does not affect ecological processes at the large population scale; and (3) individual specialisation is widespread and can significantly impact population dynamics and/or ecosystem function.

As you might have guessed by this point, I find myself in the third school of thought. There are many arguments supporting this theory, and what I believe to be very good arguments against statements 1 and 2. While I have only provided one specific named example for individual specialisation in a marine mammal, there are several documented cases of such occurrences among other marine taxa (e.g., pinnipeds14, sharks15, fish16) and a much larger number of studies for terrestrial species4. Thus, the claim that it is rare or weak, seems implausible to me.

Statement 2 is a little more complicated to tackle as it involves understanding how actions on a relatively small scale affect a whole population or even an ecosystem. For instance, consider two female sea otters living in a small coastal area where one sea otter prefers to eat turban snails and the other exclusively feeds on abalone. The sudden decline in abundance of either of these prey could lead to serious health and reproductive issues for those females. Should the low prey abundance persist, then poor health and reproduction of several females in a population that specialise on that prey item can rapidly lead to genetic loss and an overall population decline. Particularly if an individual’s or species’ home range is rather restricted or small. In the case of the sea otter, which are often touted as a keystone species due to its presence preventing sea urchin barren formation that is known to wreak havoc on kelp forests, knock-on effects of such a population decline could result in poor overall ecosystem health.

It may be easy to assume that one individual dolphin, otter, seal or whale cannot possibly make a difference to a whole population or ecosystem. This assumption strikes me as a little odd since humans are always told to ‘be the change they wish to see in the world’ and that ‘every person can make a difference’. Why then should these sentiments not be applicable to non-humans? While a gray whale may not hold a sign at a protest or run for president (actions commonly considered to cause change in the human world), perhaps the choice that a gray whale makes every day to only consume one species of zooplankton, can influence other gray whales in the area, predators from other taxa, habitat structure, other prey availability, and/or cause trophic cascades.

Through my research, I aim to elucidate whether the gray whales display some level of foraging individualisation while feeding in Port Orford, Oregon. I will use data from four years to compare tracks of individual whales with zooplankton samples collected in the area to correlate each individual’s movement patterns with prey availability. I will assess the quality of prey through bomb calorimetry and microplastic analysis of the zooplankton samples to determine energetic content and pollutant levels, respectively. This prey assessment will describe the potential effects of prey specialization on whales, which is fundamental to assessing overall population health. Individualisation can strongly affect fitness of individuals, either positively or negatively depending on several factors, which will undoubtedly have an impact at the population level.

(The videos below are examples of two different tactics we see the gray whales display while foraging along the Oregon coast in the summer months. The first video shows a whale foraging among kelp with some very acrobatic moves, while the second is of a whale employing the ‘sharking’ method where the whale is feeding benthically in such shallow depths that both the pectoral fin and the fluke stick out of the water, making the whale look like a ‘shark’.)


  1. American Psychological Association, Personality. Retrieved from:
  2. Carere C., & Locurto, C., Interaction between animal personality and animal cognition. Current Zoology, 2015. 57(4): 491-498.
  3. Gosling, S.D., From mice to men: what can we learn about personality from animal research?Psychological Bulletin, 2001. 127(1): 45-86.
  4. Bolnick, D.I., et al., The ecology of individuals: incidence and implications of individual specialisation. The American Naturalist, 2003. 161(1): 1-28.
  5. Torres, L.G., & Read, A. J., Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins (Tursiops truncatus) in Florida Bay, Florida. Marine Mammal Science, 2009. 25(4): 797-815.
  6. Gisburne, T.J., & Connor, R.C., Group size and feeding success in strand-feeding bottlenose dolphins (Tursiops truncatus) in Bull Creek, South Carolina. Marine Mammal Science, 2015. 31(3): 1252-1257.
  7. Gazda, S.K., et al., A division of labour with role specialization in group-hunting bottlenose dolphins (Tursiops truncatus) off Cedar Keys, Florida.Proceedings of the Royal Society: Biological Sciences, 2005. 272(1559): 135-140.
  8. Ford, J.K.B., et al., Dietary specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters. Canadian Journal of Zoology, 1998. 76(8): 1456-1471.
  9. Rossi-Santos, M.R., & Wedekin, L.L., Evidence of bottom contact behaviour by estuarine dolphins (Sotalia guianensis) on the Eastern Coast of Brazil.Aquatic Mammals, 2006. 32(2): 140-144.
  10. Peddemors, V.M., & Thompson, G., Beaching behaviour during shallow water feeding by humpback dolphins (Sousa plumbea). Aquatic Mammals, 1994. 20(2): 65-67.
  11. Tyack, P., Population biology, social behavior and communication in whales and dolphins. Trends in Ecology & Evolution, 1986. 1(6): 144-150.
  12. Wiley, D., et al., Underwater components of humpback whale bubble-net feeding behaviour.Behaviour, 2011. 148(5/6): 575-602.
  13. Estes, J.A., et al., Individual variation in prey selection by sea otters: patterns, causes and implications. Journal of Animal Ecology, 2003. 72(1): 144-155.
  14. Cherel, Y., et al., Stable isotopes document seasonal changes in trophic niches and winter foraging individual specialization in diving predators from the Southern Ocean. Journal of Animal Ecology, 2007. 76(4): 826-836.
  15. Matich, P., et al., Contrasting patterns of individual specialization and trophic coupling in two marine apex predators. Journal of Animal Ecology, 2010. 80(1): 294-305.
  16. Svanbäck, R., & Persson, L., Individual diet specialization, niche width and population dynamics: implications for trophic polymorphisms. Journal of Animal Ecology, 2004. 73(5): 973-982.

Oregon Sea Otter Status of Knowledge Symposium

By Dominique Kone, Masters Student in Marine Resource Management



Source: The Elakha Alliance








Source: Paul Malcolm



