Murre versus Penguin: Happy World Penguin Day!

Rachael Orben PhD, PI Seabird Oceanography Lab

Happy World Penguin Day (officially April 25th)!  I have been contemplating what to write for my tern at the GEMM lab blog. Most of my ideas were a little bit dark, but happily when I loaded my Twitter feed Saturday morning I was greeted with many beautiful photos of penguins and the hashtag #WorldPenguinDay so that inspired something more light hearted.

To be fair, it really should be Alcidae vs. Spheniscidae (scientific family names for auks and penguins). However, I have spent many months in the field studying murres (an alcid), and I find them fascinating. Soon it will be time for them to lay their eggs at colonies along the Oregon coast, including Yaquina Head. Murres have some amazing life history characteristics.

Some of the flamboyant alcid species found in the North Pacific. These species are all crevice or burrow nesters like some penguins including Magellanic, African, and little blue penguins.  

So how do murres stack up against penguins?

At first glance, murres and penguins are fairly similar. They are deep diving seabirds that forage on crustaceans and forage fish. Like murres, penguins have countershading, with black feathers on their backs and white feathers on their front. This coloring is thought to help provide camouflage when they are foraging (Cairns 1986).

There are two species of murres: common murres and thick-billed murres. Both species have a circumpolar distribution in the northern hemisphere with thick-billed murres nesting a colonies in the Arctic and common murres nesting in more temperate latitudes as far south as the central California coast. Their distributions overlap in the subarctic where they often share colonies (Irons et al. 2008).  


I am under the impression that one of the reasons people love penguins so much is because they waddle. Murres aren’t so graceful either, but they spend much less of their time walking around since they commute between the sea and their colonies by flying. However, murres have to work harder to fly than they do to dive (Elliott et al. 2013). This is because they have high wing-loading. Essentially, they have big bodies and relatively small wings that they use for flying through air and water. Bigger wings would be better for air, but smaller wings are better for moving through water.

Thick-billed murres flying home with fish, St. Paul Island, AK. Photo R. Orben

It really gets interesting when we start comparing the diving ability of alcids and penguins. Murres are the largest alcid species, and as dive depth scales with body size, they can dive the deepest. If we control for body size, alcids dive deeper then penguins (Burger 1991)! For instance, the deepest depth recorded from a thick-billed murres is 210 meters and the deepest dive of the smallest penguin (just a few hundred grams larger then the typical murre at ~1.5 kg), the little blue penguin, is a mere 69 meters (

Colonies & Nests

Murres typically nest in colonies on cliffs, off-shore sea stacks, and occasionally low lying predator free islands. Common murres use wider ledges and nest in very close proximity to each other while thick-billed murres prefer narrow ledges. Murres don’t build nests and simply lay their eggs on the rock ledge.

Common murres on Main Colony Rock at Yaquina Head, Newport Oregon. Photo R. Orben

Penguin nesting colonies can take a variety of forms. Colonies of the “brush-tailed” penguins (chinstrap, Adélie and gentoo penguins) are found in places that are snow free for most of the summer. These colonies tend to form as a meandering collection of sub-colonies.  These species build nests out of small rocks that they diligently collect. The rocks help keep their eggs out of snow meltwater. Emperor and king penguins stand together in a group. Burrow nesting penguins like Magellanic penguins can spread their colonies out across large areas where there is suitable habitat for burrowing.

A small portion of the Adélie penguin colony at Cape Crozier, Antarctica. Photo R. Orben


Murres lay one large pyriform (pear-shaped) speckled egg that ranges in color from pale cream to brilliant turquoise. This variation allows them to recognize their own eggs (Gaston et al 1993)! The purpose of the shape of murre eggs is something that has been continually puzzled over, but the shape appears to help the blunt end stay cleaner, is stronger, and is more stable on sloping surfaces (Birkhead et al. 2017, 2018).

Predated thick-billed murre eggs collected at the top of the cliffs on St. George Island, AK. Photo R. Orben

In comparison, penguin eggs don’t look that remarkable. Many penguin species lay two eggs (e.g. Adélie, chinstrap, rockhopper, gentoo), but king penguins and emperor penguins will just lay one, incubating it on top of their feet. The first egg that macaroni penguins lay is 55-75% smaller than their second egg, potentially due to constraints imposed by migration (Crossin et al. 2010).


Seabirds are not generally known for their melodious songs, but they are still an important part of their social lives. For this comparison I recommend an exploration of the Cornell Lab of Ornithology’s Macaulay Library. Start with the murres and then explore some penguin species. Recently it was discovered that penguins make short noises underwater (Thiebault 2019). Perhaps murres do as well.

If you are interesting a hearing a seabird that can sing, search for Light Mantled Sooty Albatross.

Parent-Offspring Relationship

Murres bring whole fish back to the colony to feed their chick. One fish for each trip. Murre chicks fledge before their flight feathers are fully grown. They jump from the cliffs and glide down to the ocean (hopefully) where they are joined by their male parent. Then the pair paddle out to find good foraging grounds. The male parent needs to feed the growing chick frequently and by bringing the chick to the food is able to meet these demands.

The male parent greets its newly fledged chick. Late evening on St. Paul Island, Alaska. Photo R. Orben

In contrast, penguins regurgitate their stomach contents to feed their offspring. They are able to carry large amounts of food this way. For instance a chinstrap penguin might bring back ~610 grams of food, almost 15% of its body weight (Miller et al. 2010). Adult penguins still have to balance their needs and the demands of their growing chicks. So the adults will leave their chicks alone once they are large enough. The chicks stand in groups known as créches to help protect them against predators like skuas.


Feather molt is an important part of all birds’ life histories. Feathers don’t last forever and need to be replaced. Both murres and penguins have unique strategies for replacing their feathers. For any flighted bird, replacing primary feathers is especially important. Murres become flightless during molt, which happens in the fall (Birkhead & Taylor 1977). This is actually thought to help their diving as with smaller wings they should be able to fly underwater more easily (Thompson et al. 1998). They replace their body feathers gradually to maintain waterproofing and warmth.

Penguins have solved this problem in another way. Instead of gradually replacing their feathers they undergo a “catastrophic molt” and replace all their feathers at once. Penguins need to be out of the water during this time and will fast, so it is advantageous to quickly grow a new coat of feathers. They too molt after their chicks are fledged.

I will let you decide which seabirds you find most fascinating, because really I find them all amazing and in need of our continued protection.  Thanks for reading!


Birkhead TR, Taylor AM (1977) Moult of the Guillemot Uria aalge. Ibis 119:80–85

Birkhead TR, Thompson JE, Jackson D, Biggins JD (2017) The point of a Guillemot’s egg. Ibis 159:255–265

Burger, A. E. (1991). Maximum diving depths and underwater foraging in alcids and penguins. In Studies of High-Latitude Seabirds. 1. Behavioural, Energetic and Oceanographic Aspects of Seabird Feeding Ecology (ed. W. A. Montevecchi and A. J. Gaston), pp. 9-15. Canada: Canadian Wildlife Service Occasional Paper.

Crossin GT, Trathan PN, Phillips RA, Dawson A, Le Bouard F, Williams TD (2010) A Carryover Effect of Migration Underlies Individual Variation in Reproductive Readiness and Extreme Egg Size Dimorphism in Macaroni Penguins. Am Nat 176:357–366

Elliott KH, Ricklefs RE, Gaston AJ, Hatch SA, John R Speakmane F, Davoren GK (2013) High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins. PNAS:9380–9384

Irons DB, Anker-Nilssen T, Gaston AJ, Byrd GV, Falk K, Gilchrist G, Hario M, Hjernquist M, Krasnov YV, Mosbech A, Olsen B, Petersen A, Reid JB, Robertson GJ, Strøm H, Wohl KD (2008) Fluctuations in circumpolar seabird populations linked to climate oscillations. Global Change Biology 14:1455–1463

Miller AK, Kappes MA, Trivelpiece SG, Trivelpiece WZ (2010) Foraging-Niche Separation of Breeding Gentoo and Chinstrap Penguins, South Shetland Islands, Antarctica. The Condor 112:683–695

Thiebault A (2019) First evidence of underwater vocalizations in hunting penguins. PeerJ:1–16

Thompson CW, Wilson ML, Melvin EF, Pierce DJ (1998) An unusual sequence of flight-feather molt in Common Murres and its evolutionary implications. The Auk 115:653–669

Snacks at the surface: New GEMM Lab publication reveals insights into blue whale surface foraging through drone observations and prey data

By Dawn Barlow, PhD student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

As the largest animals on the planet, blue whales have massive prey requirements to meet energy demands. Despite their enormity, blue whales feed on a tiny but energy-rich prey source: krill. Furthermore, they are air-breathing mammals searching for aggregations of prey in the expansive and deep ocean, and must therefore budget breath-holding and oxygen consumption, the travel time it takes to reach prey patches at depth, the physiological constraints of diving, and the necessary recuperation time at the surface. Additionally, blue whales employ an energetically demanding foraging strategy known as lunge feeding, which is only efficient if they can locate and target dense prey aggregations that compensate for the energetic costs of diving and lunging. In our recent paper, published today in PeerJ, we examine how blue whales in New Zealand optimize their energy use through preferentially feeding on dense krill aggregations near the water’s surface.

Figure 1. A blue whale lunges on a dense aggregation of krill at the surface. Note the krill jumping away from the mouth of the onrushing whale. UAS piloted by Todd Chandler.
Figure 2. Survey tracklines in 2017 in the South Taranaki Bight (STB) with locations of blue whale sightings, and where surface lunge feeding was observed, denoted. Inset map shows location of the STB within New Zealand. Figure reprinted from Torres et al. 2020.

To understand how predators such as blue whales optimize foraging strategies, knowledge of predator behavior and prey distribution is needed. In 2017, we surveyed for blue whales in New Zealand’s South Taranaki Bight region (STB, Fig. 2) while simultaneously collecting prey distribution data using an echosounder, which allowed us to identify the location, depth, and density of krill aggregations throughout the region. When blue whales were located, we observed their behavior from the research vessel, recorded their dive times, and used an unmanned aerial system (UAS; “drone”) to assess their body condition and behavior.

Much of what is known about blue whale foraging behavior and energetics comes from extensive studies off the coast of California, USA using accelerometer tags to track fine-scale kinematics (i.e., body movements) of the whales. In the California Current, the krill species targeted by blue whales are denser at depth, and therefore blue whales regularly dive to depths of 300 meters to lunge on the most energy-rich prey aggregations. However, given the reduced energetic costs of feeding closer to the surface, optimal foraging theory predicts that blue whales should only forage at depth when the energetic gain outweighs the cost. In New Zealand, we found that blue whales foraged where krill aggregations were relatively shallow and dense compared to the availability of krill across the whole study area (Fig. 3). Their dive times were quite short (~2.5 minutes, compared to ~10 minutes in California), and became even shorter in locations where foraging behavior and surface lunge feeding were observed.

Figure 3. Density contours comparing the depth and density (Sv) of krill aggregations at blue whale foraging sightings (red shading) and in absence of blue whales (gray shading). Density contours: 25% = darkest shade, 755 = medium shade, 95% = light shade. Blue circles indicate krill aggregations detected within 2 km of the sighting of the UAS filmed surface foraging whale analyzed in this study. Figure reprinted from Torres et al. 2020.
Figure 4. Kinematics of a blue whale foraging dive derived from a suction cup tag. Upper panel shows the dive profile (yellow line), with lunges highlighted (green circles), superimposed on a prey field map showing qualitative changes in krill density (white, low; blue, medium; red, high). The lower panels show the detailed kinematics during lunges at depth. Here, the dive profile is shown by a black line. The orange line shows fluking strokes derived from the accelerometer data, the green line represents speed estimated from flow noise, and the grey circles indicate the speed calculated from the vertical velocity of the body divided by the sine of the body pitch angle, which is shown by the red line. Figure and caption reprinted from Goldbogen et al. 2011.

Describing whale foraging behavior and prey in the surface waters has been difficult due to logistical limitations of conventional data collection methods, such as challenges inferring surface behavior from tag data and quantifying echosounder backscatter data in surface waters. To compliment these existing methods and fill the knowledge gap surrounding surface behavior, we highlight the utility of a different technological tool: UAS. By analyzing video footage of a surface lunge feeding sequence, we obtained estimates of the whale’s speed, acceleration, roll angle, and head inclination, producing a figure comparable to what is typically obtained from accelerometer tag data (Fig. 4, Fig. 5). Furthermore, the aerial perspective provided by the UAS provides an unprecedented look at predator-prey interactions between blue whales and krill. As the whale approaches the krill patch, she first observes the patch with her right eye, then turns and lines up her attack angle to engulf almost the entire prey patch through her lunge. Furthermore, we can pinpoint the moment when the krill recognize the impending danger of the oncoming predator—at a distance of 2 meters, and 0.8 seconds before the whale strikes the patch, the krill show a flee response where they leap away from the whale’s mouth (see video, below).

Figure 5. Body kinematics during blue whale surface lunge feeding event derived from Unmanned Aerial Systems (UAS) image analysis. (A) Mean head inclination and roll (with CV in shaded areas), (B) relative speed and acceleration, and (C) distance from the tip of the whale’s rostrum to the nearest edge of krill patch. Blue line on plots indicate when krill first respond to the predation event, and the purple dashed lines indicate strike at time = 0. The orange lines indicate the time at which the whale’s gape is widest, head inclination is maximum, and deceleration is greatest. Figure reprinted from Torres et al. 2020

In this study, we demonstrate that surface waters provide important foraging opportunities and play a key role in the ecology of New Zealand blue whales. The use of UAS technology could be a valuable and complimentary tool to other technological approaches, such as tagging, to gain a comprehensive understanding of foraging behavior in whales.

To see the spectacle of a blue whale surface lunge feeding, we invite you to take a look at the video footage, below:

The publication is led by GEMM Lab Principal Investigator Dr. Leigh Torres. I led the prey data analysis portion of the study, and co-authors include our drone pilot extraordinaire Todd Chandler and UAS analysis guru Dr. Jonathan Burnett. We are grateful to all who assisted with fieldwork and data collection, including Kristin Hodge, Callum Lilley, Mike Ogle, and the crew of the R/V Star Keys (Western Workboats, Ltd.). Funding for this research was provided by The Aotearoa Foundation, The New Zealand Department of Conservation, The Marine Mammal Institute at Oregon State University, Greenpeace New Zealand, OceanCare, Kiwis Against Seabed Mining, The International Fund for Animal Welfare, and The Thorpe Foundation.

Read Oregon State University’s press release about the publication here.

Whale blow: good for more than spotting whales

Clara Bird, Masters Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

Whale blow, the puff of air mixed with moisture that a whale releases when it comes to the surface, is a famously thrilling indicator of the presence of a whale. From shore, spotting whale blow brings the excitement of knowing that there are whales nearby. During boat-based field work, seeing or hearing blow brings the rush of adrenaline meaning that it’s game time. Whale blow can also be used to identify different species of whales, for example gray whale blow is heart shaped (Figure 1). However, whale blow can be used for more than just spotting and identifying whales. We can use the time between blows to study energetics.

Figure 1. Gray whale blow is often heart shaped (when there is very little wind). Source:

A blow interval is the time between consecutive blows when a whale is at the surface (Stelle, Megill, and Kinzel 2008). These are also known as short breath holds, whereas long breath holds are times between surfacings (Sumich 1983).  Sumich (1983) hypothesized that short breath holds lead to efficient rates of oxygen use. The body uses oxygen to create energy, so “efficient rate of oxygen use” means that longer breath holds do not use much more oxygen and subsequently do not produce more energy.  Surfacings, during which short blow intervals occur, are often thought of as recovery periods for whales. Think of it this way, when you sprint, immediately afterwards you typically need to take a break to just breathe and recover.

We hypothesize that we can use blow intervals as a measure of how strenuous an activity is; shorter blow intervals may indicate that an activity is more energetically demanding (Wursig, Wells, and Croll 1986). Let’s go back to the sprinting analogy and compare the energetic demands of walking and running. Imagine I asked you to walk for five minutes, stop and measure the time between each breath, and then run for five minutes and do the same; after running, you would likely breathe more heavily and take more breaths with less time between them. This result indicates that running is more demanding, which we already know because we can do other experiments with humans to study metabolic rate and related metrics. In the case of gray whales, we cannot do experiments in the same way, but we can use the same analogy. Several studies have examined how blow intervals differ between travelling and foraging.

Wursig, Wells, and Croll (1986) measured blow interval, surfacing time, and estimated dive depth and duration of gray whales in Alaska from a boat during the foraging season. They found that blow intervals were shorter during feeding. They also found that the number of blows per surfacing increased with increasing depth. Overall these findings suggest that during the foraging season, feeding is more strenuous than other behaviors and that deeper dives may be more physiologically stressful.

Stelle, Megill, and Kinzel (2008) studied gray whales foraging off of British Columbia, Canada. They found shorter blow intervals during foraging, intermediate blow intervals during searching, and longer blow intervals during travelling. Interestingly, within feeding behaviors, they found a difference between whales feeding on mysids (krill-like animals that swim in the water column) and whales feeding benthically on amphipods. They found that whales feeding on mysids made more frequent but shorter dives with short blow intervals at surface, while whales feeding benthically had longer dives with longer blow intervals. They hypothesized that this difference in surfacing pattern is because mysids might scatter when disturbed, so gray whales surface more often to allow the mysids swarm to reform. These studies inspired me to start investigating these same questions with my drone video data.

As I review the drone footage and code the behaviors I also mark the time of each blow. I’ve done some initial video coding and using this data I have started to look into differences in blow intervals. As it turns out, we see a similar difference in blow interval relative to behavior state in our data: whales that are foraging have shorter blow intervals than when traveling (Figure 2). It is encouraging to see that our data shows similar patterns.

Figure 2. Boxplot of mean blow interval per sighting of foraging whales and travelling whales.

Next, I would like to examine how blow intervals differ between foraging tactics. A significant part of my thesis is dedicated to studying specific foraging tactics. The perspective from the drone allows us to identify behaviors in greater detail than studies from shore or boat (Torres et al. 2018), allowing us to dig into the differences between the different foraging behaviors. The purpose of foraging is to gain energy. However, this gain is a net gain. To understand the different energetic “values” of each tactic we need to understand the cost of each behavior, i.e. how much energy is required to perform the behavior. Given previous studies, maybe blow intervals could help us measure this cost or at least compare the energetic demands of the behaviors relative to each other. Furthermore, because different behaviors are likely associated with different prey types (Dunham and Duffus 2001), we also need to understand the different energetic gains of each prey type (this is something that Lisa is studying right now, check out the COZI project to learn more). By understanding both of these components – the gains and costs – we can understand the energetic tradeoffs of the different foraging tactics.

Another interesting component to this energetic balance is a whale’s health and body condition. If a whale is in poor health, can it afford the energetic costs of certain behaviors? If whales in poor body condition engage in different behavior patterns than whales in good body condition, are these patterns explained by the energetic costs of the different foraging behaviors? All together this line of investigation is leading to an understanding of why a whale may choose to use different foraging behaviors in different situations. We may never get the full picture; however, I find it really exciting that something as simple and non-invasive as measuring the time between breaths can contribute such a valuable data stream to this project.


Dunham, Jason S., and David A. Duffus. 2001. “Foraging Patterns of Gray Whales in Central Clayoquot Sound, British Columbia, Canada.” Marine Ecology Progress Series 223 (November): 299–310.

Stelle, Lei Lani, William M. Megill, and Michelle R. Kinzel. 2008. “Activity Budget and Diving Behavior of Gray Whales (Eschrichtius Robustus) in Feeding Grounds off Coastal British Columbia.” Marine Mammal Science 24 (3): 462–78.

Sumich, James L. 1983. “Swimming Velocities, Breathing Patterns, and Estimated Costs of Locomotion in Migrating Gray Whales, Eschrichtius Robustus.” Canadian Journal of Zoology 61 (3): 647–52.

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).

Wursig, B., R. S. Wells, and D. A. Croll. 1986. “Behavior of Gray Whales Summering near St. Lawrence Island, Bering Sea.” Canadian Journal of Zoology 64 (3): 611–21.

Humans Hide and Wildlife Thrive: Human-mediated ecosystem changes during a pandemic

By: Alexa Kownacki, Ph.D. Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

We live in an interesting time. Many of us academic scientists sit in the confines of our homes, reading scientific papers, analyzing years-worth of data, working through a years-worth of house projects, or simply watching Netflix. While we are confined to a much smaller area, wildlife is not.  

During this challenging situation we have unique opportunities to study what happens when people are not outside for recreation. All of us who feel trapped inside our homes are not only saving human lives, we are changing ecosystems. Humans are constantly molding our ecosystems on fine and grand scales, from xeriscaping our lawns with native, drought-resistant plants to developing large plots of land for new homes. We manipulate nature, for better or for worse.

So, what happens when we change our behavior? Rather than driving, we’re gardening, instead of playing at parks, we’re playing board games at our kitchen tables; we as a society are completely changing our habitat-use patterns. When any top predator changes its habitat-use, switches niches, or drastically changes its behaviors, there are top-down ecosystem effects. When one species changes its behavior, there are major downstream impacts on predation, foraging, diet, and habitat use. For example, when bluegill sunfish underwent large shifts in both diet and habitat, major predator-mediated habitat use changes in other species occurred (Mittelbach 1986). There are multiple studies describing the impacts of human-mediated drivers on ecosystems worldwide. In coastal environments, anthropogenic activities, specifically shipping, industry, and urban development, dramatically change both the coastal and marine ecosystems (Mead et al. 2013).

The highly developed coastline along Los Angeles, CA is a prime example of urban development. (Image source: LA Magazine.)

By far the most pronounced example of how an international halt on travel can alter ecosystems comes from the tragic terrorist attacks on September 11, 2001. Prior to this current, viral pandemic, the events following 9/11 were the first time that nearly all major transit stopped in the USA—including airplanes and major shipping traffic. This halt created a unique opportunity to study some of the secondary impacts, such as a reduction in shipping traffic noise, on cetaceans. Following 9/11, there was a six decibel decrease in underwater noise that co-occurred with a decrease in stress hormones of endangered North Atlantic right whales (Rolland et al. 2012). When I first read about this study, my first thought was “leave it to scientists to make the best out of a terrible situation.” Truly, learning from nature, even in the darkest of days, is an incredible skillset. Research like this inspires me to ask questions about what changes are happening in ecosystems now because of recent events. For example, the entire port of San Diego, its beaches and bays, are closed for all recreational activity and I wonder how this reduction in traffic is similar to the post-9/11 study but on bottlenose dolphins, gray whales, and pinnipeds that are coast-associated. Are urban and suburban neighborhoods slowly becoming more rural and making space for wildlife again?

My dad lives in a suburban neighborhood of San Diego, CA. In the past few weeks, his “Ring doorbell camera” captured a bobcat walking along the raised brick path multiple times. (Media source: Eric Kownacki)

There is increasing news coverage on wild animals “taking over” cities. Dr. Leila Lemos touched on this earlier with her blog post centering on how academics are changing their means of teaching, conferencing, and learning. There are photos of wild goats running through the streets of Wales, UK, coyotes roaming the streets of San Francisco, CA, USA, monkeys swarming the streets in Thailand, pumas wandering the streets of Santiago, Chile, and Sika deer peering into empty restaurants in Nara, Japan (Colarossi 2020). In reality, this wildlife was likely part of the ecosystem prior to the formation of these cities but was forced out of the more urban centers. As we sit in our homes, rather than looking bleakly onto empty streets, we can search for wildlife, create a backyard birding competition with your friends, guess which flowers will bloom first, and ask questions of our changing ecosystems.

Coyote at a park in northern California with the San Francisco Golden Gate Bridge in the background. (Image source: u/beccatravels via Reddit)


Colarossi, Natalie. “Photos Show Wild Animals Roaming Empty Streets as Coronavirus Lockdowns Keep Humans Inside.” Insider, Insider, 2 Apr. 2020,

Mead, A., Griffiths, C.L., Branch, G.M., McQuaid, C.D., Blamey, L.K., Bolton, J.J., Anderson, R.J., Dufois, F., Rouault, M., Froneman, P.W. and Whitfield, A.K., 2013. Human-mediated drivers of change—impacts on coastal ecosystems and marine biota of South Africa. African Journal of Marine Science35(3), pp.403-425.

Mittelbach, Gary. 1986. Predator-mediated habitat use: some consequences for species interactions. Environ Biol Fish 16, 159–169.

Rolland, R.M., Parks, S.E., Hunt, K.E., Castellote, M., Corkeron, P.J., Nowacek, D.P., Wasser, S.K. and Kraus, S.D., 2012. Evidence that ship noise increases stress in right whales. Proceedings of the Royal Society B: Biological Sciences279(1737), pp.2363-2368.