Tag: behavior

Every breath [a whale] takes: How and why we study cetacean respiration

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

We need energy to function and survive. For animals in the wild who may have limited food availability, knowing how they spend their energy is a critical question for many scientists because it fundamentally informs how we understand their decisions about where they go and what they do. The entire field of foraging theory is founded on the concept that animals optimize their ratio of energy in and energy out so that they have enough energy to survive, reproduce (pass on their genes), watch out for threats, if need be, and rest. And, if we understand an animal’s ‘typical’ energy budget, we can then try to predict how disturbance or environmental change will affect their actual energy budgets as a consequence of that change. But how do we measure energy expenditure?

The most commonly measured energy currency is oxygen. Since our cells use oxygen to produce energy (this is why we need oxygen to live), we can measure oxygen consumption as a metric of energy expenditure. The more oxygen we consume, the more energy we’re expending. In ideal lab settings, oxygen consumption can be accurately measured by placing the subject in a chamber where the oxygen flow can be controlled (Speakman, 1999). However, you can probably see how that approach is problematic for measuring oxygen consumption in most large free-living animals, especially cetaceans. It isn’t exactly feasible to put a whale in a box.

Image 1. A great tit in a metabolic chamber. Figure 1 from Broggi et al., 2009

Fortunately, a tool called a spirometer was developed to measure oxygen consumption in restrained cetaceans. A spirometer is a device that can be placed over the blowhole(s) of an individual to accurately measure the amount of air that is exhaled and inhaled (Figure 1).  For trained cetaceans in captivity (e.g., dolphins), spirometers can be used to quantify how respiration changes after the animal performs certain behaviors (Fahlman et al., 2019). The breathing patterns of diving mammals are particularly interesting because they cannot breathe during most of their exercise (energy expenditure) as they are underwater. So, their breathing patterns after a dive tell us a lot about how much energy they spent during that dive. For example, Fahlman et al. (2019) used spirometer data from dolphins in captivity to study how their breathing patterns changed while recovering from dives of different durations. Interestingly, they found that after longer dives, dolphins took larger breaths (i.e., inhaled more air) while recovering but did not change the number of breaths. This finding is particularly relevant to the work we are conducting in the GEMM lab, where we utilize breathing patterns to quantify the energy expenditure of cetaceans in the wild, where spirometers cannot be used.

Figure 1. Figure 1 from Sumich et al. (2023). Left: a spirometer being held over the blow holes of JJ, a gray whale calf at sea world in 1997; one of the rare times that a large baleen whale was in captivity and available for these measurements. Right: example of a plot created using the data from a spirometer over JJ’s blow holes. The duration of a “blow” (exhale followed by immediate inhale) is on the x-axis, the flow rate (in liters per second) is on the y-axis. The positive curve during the exhale shows that the whale strongly exhales a lot of air very quickly, then the negative curve shows the whale inhaling a lot of air very quickly.

In a previous blog, I described how inter-breath intervals (the time between consecutive blows) are useful for estimating energy expenditure in free-living cetaceans. Essentially, a shorter interval indicates that the whale was just engaged in an energetically demanding activity. When you’re recovering from a sprint, you breathe faster (i.e., with shorter inter-breath intervals), than when you’re recovering from a walk. However, a big assumption in using inter-breath intervals as a proxy for energy expenditure is that every breath is equal. But as Fahlman et al. emphasize in their 2016 paper, every blow is not equal (Fahlman et al., 2016). In addition to varying the time between breaths, an animal can vary the intensity of each breath (e.g., Fahlman et al., 2019), the duration of each breath (Sumich et al., 2023), the number of breaths, and even the expansion of their nostrils (Nazario et al., 2022; check out this blog for more).

Image 2. Gray whale blow. Source: https://www.lajollalight.com/sdljl-natural-la-jolla-winter-wildlife-2015jan08-story.html

Altogether, this means that it’s important to measure every breath and that no one metric tells the complete story. This also means my research question focused on comparing the energetic costs of different tactics is more complicated than I originally thought. If we go back to the first blog I wrote on this topic, I was planning ons only using inter-breath intervals to estimate energy expenditure. Fast forward four years, with all my new knowledge gained on respiration variability, I’ve modified my plan and now I’m working to first understand how all these different metrics of breathing relate to each other. Then, I’ll compare how breathing varies between different foraging tactics, which is an important follow up to my questions around individual specialization of foraging tactics. If different whales are using different foraging behaviors, does that mean they’re spending different amounts of energy? If so, are certain behaviors more advantageous than others? Of course, these answers are incomplete without understanding the prey the whales are eating, but that’s something that PhD student Nat Chazal is working to understand (check out her recent blog here).  For now, I’m working on bringing integrating all the measures of breathing, then we will start putting the story together and finding some answers to our pressing questions. 

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References

Broggi, J., Hohtola, E., Koivula, K., Orell, M., & Nilsson, J. (2009). Long‐term repeatability of winter basal metabolic rate and mass in a wild passerine. Functional Ecology, 23(4), 768–773. https://doi.org/10.1111/j.1365-2435.2009.01561.x

Fahlman, A., Brodsky, M., Miedler, S., Dennison, S., Ivančić, M., Levine, G., Rocho-Levine, J., Manley, M., Rocabert, J., & Borque-Espinosa, A. (2019). Ventilation and gas exchange before and after voluntary static surface breath-holds in clinically healthy bottlenose dolphins, Tursiops truncatus. Journal of Experimental Biology, 222(5), jeb192211. https://doi.org/10.1242/jeb.192211

Fahlman, A., van der Hoop, J., Moore, M. J., Levine, G., Rocho-Levine, J., & Brodsky, M. (2016). Estimating energetics in cetaceans from respiratory frequency: Why we need to understand physiology. Biology Open,5(4), 436–442. https://doi.org/10.1242/bio.017251

Nazario, E. C., Cade, D. E., Bierlich, K. C., Czapanskiy, M. F., Goldbogen, J. A., Kahane-Rapport, S. R., Hoop, J. M. van der, Luis, M. T. S., & Friedlaender, A. S. (2022). Baleen whale inhalation variability revealed using animal-borne video tags. PeerJ, 10, e13724. https://doi.org/10.7717/peerj.13724

Speakman, J. R. (1999). The Cost of Living: Field Metabolic Rates of Small Mammals. In A. H. Fitter & D. G. Raffaelli (Eds.), Advances in Ecological Research (Vol. 30, pp. 177–297). Academic Press. https://doi.org/10.1016/S0065-2504(08)60019-7

Sumich, J. L., Albertson, R., Torres, L. G., Bird, C. N., Bierlich, K. C., & Harris, C. (2023). Using audio and UAS-based video for estimating tidal lung volumes of resting and active adult gray whales (Eschrichtius robustus). Marine Mammal Science, 1(8). https://doi.org/10.1111/mms.13081

That’s so Real: Adult Beginners, Serial Podcast(s), and a whole lotta of Baja Gray Whale Video Analysis.

Celest Sorrentino, Research Technician, Geospatial Ecology of Marine Megafauna Lab

Hello again GEMM Lab family. I write to you exactly a year after (okay maybe 361 days after but who’s counting…) from my previous blog post describing my 2022 summer working in the GEMM Lab as an NSF REU intern. Since then, so much has changed, and I can’t wait to fill you in on it.

In June I walked across the commencement stage at UC Santa Barbara, earning my BS in Ecology, Evolution, and Marine Biology and my minor in Italian language. A week later, I packed my bags and headed straight back to the lukewarm beaches of Newport, Oregon as a Research Technician in the GEMM Lab. I am incredibly fortunate to have been invited back to the OSU Marine Mammal Institute to lend a hand analyzing drone footage of gray whales collected back in March 2023 when Leigh and Clara went down to Baja California, as mentioned previously in Clara’s blog. 

Fig. 1. View from the top! (of the bridge at Yaquina Bay Bridge in Newport, OR)

During my first meeting with Clara at the beginning of the summer we discussed that a primary goal of my position was to process all the drone footage collected in Baja so that the generated video clips could be later used in other analytical software such as BORIS and SLEAP A.I. Given my previous internships and past summer project, this video processing is familiar to me. My initial thoughts were:

Sweet! Watch drone footage, pop in some podcasts, note down when I see whales, let’s do this!*

Like any overly eager 23-year-old, I might have mentally cracked open a Celsius and kicked my feet up too soon. We added another layer to the goal: develop an ethogram – which requires me to identify and define the behaviors that the gray whales appear to be demonstrating within the videos (more on ethogram development in Clara’s previous blog.) This made me nervous. 

I don’t have any experience with behavior. How do I tell what is a real behavior or if the whale is just existing? What if I’m wrong and ruin the project? What if I totally mess this up?

Naturally, as any sane person, to resolve these thoughts I took to the Reddit search bar: “How to do a job you’ve never done before.” No dice. 

I pushed these thoughts aside and decided to just start the video analysis process. Clara provided me with the ethogram she is developing during her PhD as a point of reference (based on the published gray whale ethogram in Torres et al. 2018), I was surrounded by an insanely supportive lab, and I could Google anything at my fingertips. Fast-forward 6 weeks later: I had analyzed 128 drone videos of adult gray whales as well as mother-calf pairs, and developed an ethogram describing, 26 behaviors**. I named one of my favorite behaviors  a “Twirl” to describe when a gray whale lifts their head out of the water and performs a 360 turn. Reminds me of times when as a kid, sometimes all you really needed is a good spin!

Now I was ready to start a productive, open conversation with Leigh and Clara about this ethogram and my work. However, even walking up to that last meeting, remnants of those daunting, doubtful early summer thoughts persisted. Even after I double checked all the definitions I wrote, rewatched all videos with said behaviors, and had something to show for my work. What gives Brain?

A few days ago, as I sat on my family’s living room couch with my two younger sisters, Baylie and Cassey, Baylie wanted to watch some TikToks with me. One video that came up was of a group of adults taking a beginner dance class, having so much fun and radiating joy. The caption read, Being a beginner as an adult is such a fun and wild thing. Baylie and I watched the video at least 10x, repeating to each other phrases like, “Wow!” and “They’re so cool.” That caption and video has been on my mind since: 

Being a beginner as an adult is such a fun and wild thing.

Being a beginner as an adult is also scary. 

Having just graduated, I can no longer say I am undergraduate student. Now, I am a young adult. This was my first research technician job, as an adult. Don’t adults usually have everything figured out? Can adults be beginners too?

Yes. In fact, we’re beginners more than we realize. 

  • I was a beginner cooking my mother’s turkey recipe 3 years ago for my housemates during the pandemic (Even after having her on Facetime, I still managed to broil it a little too long.) 
  • I was a beginner driver 5 years ago in a rickety Jeep driving myself to school (Now, since I’ve been back home, I’ve been driving my little sisters to school.)
  • I was a beginner NSF REU intern just a year ago. (This summer I was the alumni on the panel for the current NSF REU interns at Hatfield.)
  • I was a beginner science communicator presenting my NSF REU project at Hatfield last summer. (This summer, I presented my research at the Animal Behavior Society Conference.) 
Fig 2A. Group Pic with the LABIRINTO Lab and GEMM Lab at the ABS Portland Conference!
Fig 2B. Clara Bird (left), Dr. Leigh Torres (middle), and I (right) at the ABS Portland Conference. 

I now recognize that during my time identifying and defining behaviors of gray whales in videos made me take on the seat of a “beginner video and behavioral analyst”. I could not rely on the automated computer vision lens I gained from previous internships, which felt familiar and secure. 

 Instead, I had to allow myself to be creative. Dig into the unfamiliar in an effort to complete a task or job I had never done before. Allowing myself to be imperfect, make mistakes, meanwhile unconsciously building a new skill. 

This is what makes being a beginner as an adult such a fun thing. 

I don’t think being a beginner is a wild thing, although it can definitely make you feel a wild range of emotions. Being a beginner means you’re allowing yourself to try something new. Being a beginner means you’re allowing yourself the chance to learn.

Whether you’re an adult beginner as you enter your 30s, adult beginner as you enter parenthood, adult beginner grabbing a drink with friends after a long day in lab, adult beginner as a dancer, or like me, a beginner of leaving behind my college student persona and entering a new identity of adulthood, being a beginner as an adult is such a fun and normal thing.

I am not sure what will be next, but I hope to write to you all again from this blog a year from now, as an adult beginner as a grad student in the GEMM Lab. For anyone approaching the question of “What’s next”, I encourage you to read “Never a straight Path” by GEMM Lab MSc alum Florence Sullivan, a blog that has brought me such solace in my new adult journey and advice that never gets old.

Being a beginner—that, is so real. 

Fig 3A. Kayaking as an adult beginner of the Port Orford Field Team!
Fig 3B “See you soon:” Wolftree evenings with the lab.
Fig 3C. GEMM Lab first BeReal!

*I listened to way too many podcasts to list them all, but I will include two that have been a GEMM Lab “gem” —-thanks to Lisa and Clara for looping me in and now, looping you in!)

**(subject to change)

References

Torres LG, Nieukirk SL, Lemos L, Chandler TE (2018) Drone Up! Quantifying Whale Behavior From a New Perspective Improves Observational Capacity. Front Mar Sci 510.3389/fmars.2018.00319

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How do we study the impact of whale watching?

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

Since its start, the GEMM Lab has been interested in the effect of vessel disturbance on whales. From former student Florence’s masters project to Leila’s PhD work, this research has shown that gray whales on their foraging grounds have a behavioral response to vessel presence (Sullivan & Torres, 2018) and a physiological response to vessel noise (Lemos et al., 2022). Presently, our GRANITE project is continuing to investigate the effect of ambient noise on gray whales, with an emphasis on understanding how these effects might scale up to impact the population as a whole (Image 1).

To date, all this work has been focused on gray whales feeding off the coast of Oregon, but I’m excited to share that this is about to change! In just a few weeks, Leigh and I will be heading south for a pilot study looking at the effects of whale watching vessels on gray whale mom/calf pairs in the nursing lagoons of Baja California, Mexico.

Image 1. Infographic for the GRANITE project. Credit: Carrie Ekeroth

We are collaborating with a Fernanda Urrutia Osorio, a PhD candidate at Scripps Institute of Oceanography, to spend a week conducting fieldwork in one of the nursing lagoons. For this project we will be collecting drone footage of mom/calf pairs in both the presence and absence of whale watching vessels. Our goal is to see if we detect any differences in behavior when there are vessels around versus when there are not. Tourism regulations only allow the whale watching vessels to be on the water during specific hours, so we are hoping to use this regulated pattern of vessel presence and absence as a sort of experiment.

Image 2. A mom and calf pair.  NOAA/NMFS permit #21678.

The lagoons are a crucial place for mom/calf pairs, this is where calves nurse and grow before migration, and nursing is energetically costly for moms. So, it is important to study disturbance responses in this habitat since any change in behavior caused by vessels could affect both the calf’s energy intake and the mom’s energy expenditure. While this hasn’t yet been investigated for gray whales in the lagoons, similar studies have been carried out on other species in their nursing grounds.

Video 1. Footage of “likely nursing” behavior. NOAA/NMFS permit #21678.

We can use these past studies as blueprints for both data collection and processing. Disturbance studies such as these look for a wide variety of behavioral responses. These include (1) changes in activity budgets, meaning a change in the proportion of time spent in a behavior state, (2) changes in respiration rate, which would reflect a change in energy expenditure, (3) changes in path, which would indicate avoidance, (4) changes in inter-individual distance, and (5) changes in vocalizations. While it’s not necessarily possible to record all of these responses, a meta-analysis of research on the impact of whale watching vessels found that the most common responses were increases in the proportion of time spent travelling (a change in activity budget) and increased deviation in path, indicating an avoidance response (Senigaglia et al., 2016).

One of the key phrases in all these possible behavioral responses is “change in ___”. Without control data collected in the absence of whale watching vessels, it impossible to detect a difference. Some studies have conducted controlled exposures, using approaches with the research vessel as proxies for the whale watchers (Arranz et al., 2021; Sprogis et al., 2020), while others use the whale watching operators’ daily schedule and plan their data collection schedule around that (Sprogis et al., 2023). Just as ours will, all these studies collected data using drones to record whale behavior and made sure to collect footage before, during, and after exposure to the vessel(s).

One study focused on humpback mom/calf pairs found a decrease in the proportion of time spent resting and an increase in both respiration rate and swim speed during the exposure (Sprogis et al., 2020). Similarly, a study focused on short-finned pilot whale mom/calf pairs found a decrease in the mom’s resting time and the calf’s nursing time (Arranz et al., 2021). And, Sprogis et al.’s  study of Southern right whales found a decrease in resting behavior after the exposure, suggesting that the vessels’ affect lasted past their departure (Sprogis et al., 2023, Image 3). It is interesting that while these studies found changes in different response metrics, a common trend is that all these changes suggest an increase in energy expenditure caused by the disturbance.

However, it is important to note that these studies focused on short term responses. Long term impacts have not been thoroughly estimated yet. These studies provide many valuable insights, not only into the response of whales to whale watching, but also a look at the various methods used. As we prepare for our fieldwork, it’s useful to learn how other researchers have approached similar projects.

Image 3. Visual ethogram from Sprogis et al. 2023. This shows all the behaviors they identified from the footage.

I want to note that I don’t write this blog intending to condemn whale watching. I fully appreciate that offering the opportunity to view and interact with these incredible creatures is valuable. After all, it is one of the best parts of my job. But hopefully these disturbance studies can inform better regulations, such as minimum approach distances or maximum engine noise levels.

As these studies have done, our first step will be to establish an ethogram of behaviors (our list of defined behaviors that we will identify in the footage) using our pilot data. We can also record respiration and track line data. An additional response that I’m excited to add is the distance between the mom and her calf. Former GEMM Lab NSF REU intern Celest will be rejoining us to process the footage using the AI method she developed last summer (Image 4). As described in her blog, this method tracks a mom and calf pair across the video frames, and allows us to extract the distance between them. We look forward to adding this metric to the list and seeing what we can glean from the results.

Image 4. Example of a labelled frame from SLEAP, highlighting labels: rostrum, blowhole, dorsal, dorsal-knuckle, and tail. This labels are drawn to train the software to recognize the whales in unlabelled frames.

While we are just getting started, I am excited to see what we can learn about these whales and how best to study them. Stay tuned for updates from Baja!

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References

Arranz, P., Glarou, M., & Sprogis, K. R. (2021). Decreased resting and nursing in short-finned pilot whales when exposed to louder petrol engine noise of a hybrid whale-watch vessel. Scientific Reports, 11(1), 21195. https://doi.org/10.1038/s41598-021-00487-0

Lemos, L. S., Haxel, J. H., Olsen, A., Burnett, J. D., Smith, A., Chandler, T. E., Nieukirk, S. L., Larson, S. E., Hunt, K. E., & Torres, L. G. (2022). Effects of vessel traffic and ocean noise on gray whale stress hormones. Scientific Reports, 12(1), Article 1. https://doi.org/10.1038/s41598-022-14510-5

Senigaglia, V., Christiansen, F., Bejder, L., Gendron, D., Lundquist, D., Noren, D., Schaffar, A., Smith, J., Williams, R., Martinez, E., Stockin, K., & Lusseau, D. (2016). Meta-analyses of whale-watching impact studies: Comparisons of cetacean responses to disturbance. Marine Ecology Progress Series, 542, 251–263. https://doi.org/10.3354/meps11497

Sprogis, K. R., Holman, D., Arranz, P., & Christiansen, F. (2023). Effects of whale-watching activities on southern right whales in Encounter Bay, South Australia. Marine Policy, 150, 105525. https://doi.org/10.1016/j.marpol.2023.105525

Sprogis, K. R., Videsen, S., & Madsen, P. T. (2020). Vessel noise levels drive behavioural responses of humpback whales with implications for whale-watching. ELife, 9, e56760. https://doi.org/10.7554/eLife.56760

Sullivan, F. A., & Torres, L. G. (2018). Assessment of vessel disturbance to gray whales to inform sustainable ecotourism. Journal of Wildlife Management, 82(5), 896–905. https://doi.org/10.1002/jwmg.21462

What drives individual specialization?

Clara Bird, PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

When I wrote my first blog on individual specialization well over a year ago, I just skimmed the surface of the literature on this topic and only started to recognize the importance of studying individual specialization. The question, “is there individual specialization in the PCFG of gray whales?” is the focus of my first thesis chapter and the results will affect all my subsequent work. Therefore, the literature and concepts of individual specialization are a focus of my literature review and studies.

In my previous blog I focused on common characteristics of individuals that are similarly specialized as an underlying driver of individual specialization. While these characteristics (often attributable to age, sex, or physical traits) are important to consider, I’ve learned that the list of drivers of individual specialization is long and that many variables are dynamic. Of all the drivers I’ve learned about, competition is among the most common.

Competition is a major driver of individual specialization, and a common driver of competition is resource availability. When resource availability decreases, whether caused by increasing population density or changing environmental conditions, competition for that resource increases. As competition increases, individuals have a choice. They can choose to engage in competition, either by racing, fighting, or sharing [1], which can be costly, or they can diffuse the competition by focusing on a different resource.  This second approach would be considered niche partitioning in the prey dimension. Niche partitioning is a way for individuals to share ecological space by using different resources. Essentially, individuals can share habitat without having to engage in direct competition by pursuing different prey types [2]. 

This switch to different prey types can change the degree of individual specialization present in the population (Figure 1). But the direction of the change is not constant. If all individuals were pursuing the same prey type under low competition conditions but then switched to different alternate prey types under high competition, then individual specialization would increase (Figure 1a). This direction has been observed across a range of species including sharks [3], otters [4]–[7], dolphins [8], [9], stickleback fish [10], [11], largemouth bass [12], banded mongoose [13], fur seals [14], and baleen whales [15].

However, if individuals were pursuing different prey types under low competition conditions (maybe because of underlying differences such as age or sex) but then switched to the same alternate prey types under high competition, diet overlap would increase, and individual specialization would decrease (Figure 1b). Furthermore, an individual might not switch to an entirely new prey type but instead add prey items to its diet [16]. This diet expansion under competition would also decrease individual specialization. Fewer studies have reported this direction but it’s been found in the common bumblebee [17] and in several neotropical vertebrate species [18], [19].

Figure `1. Figure 3 from Araújo et al. 2011 [20]. Illustration of how ecological mechanisms may affect the degree of individual specialization. Arrows linking resources to individual consumers indicate resource consumption (relative thickness indicates proportional contribution). 
Horizontal arrows indicate the sign (positive or negative) of the effect on the degree of individual specialization. (a) Consumers share the same preferred resource (dark gray tangle) but have different alternative resources (white and light gray triangles). As the preferred resource becomes scarce, consumers switch to different alternatives, increasing the degree of individual specialization. (b) Alternatively, consumers have distinct preferred resources, so that as resources become scarce, individuals converge to the alternative resource (dark gray triangle), reducing diet variation.

Interestingly, its hypothesized that individual specialization driven by competition is one of the factors that facilitates the formation and existence of stable groups [21]. For example, a study on resident female dolphins in Sarasota Bay, FL, USA found that females with high spatial overlap used distinct foraging specializations [8](Figure 2). This study illustrates how partitioning prey enabled spatial and social coexistence. A study on banded mongooses reached a similar conclusion [13]. They found that specialization was highest in the biggest groups (with the most competition) and not explained by sex, age, or other inherent differences. They hypothesized that specialization increasing with competition reduced conflict and allowed the groups to remain stable. This study also highlighted the role of learning to determine an individual’s specialization.

Figure 2. A bottlenose dolphin.
Source: https://sarasotadolphin.org

Learning drives the distribution of knowledge throughout a population, which can lead to either specialization or generalization. ‘One-to-one’ learning, where one individual learns from one demonstrator, tends to promote individual specialization [21]. This form of transmission drives specialization because the individuals who learn the specialization tend to then carry on using, and eventually teaching, that specialization [6]. A common example of ‘one-to-one’ learning is vertical transmission from parent to offspring. It has been shown to transmit specializations in dolphins [22] and otters [6]. ‘One-to-one’ learning can occur outside of parent-offspring pairs; non-parent-offspring ‘one-to-one’ learning has been shown to drive specialization in banded mongooses [13](Figure 3).

However, other forms of social learning can promote more generalized foraging strategies. ‘Many-to-one’ or ‘one-to-many’ learning  can reduce the presence of specialization in species [13], [21] as can the presence of conformity in a group [23], [24].

Figure 3. A group of banded mongooses.
Source: http://socialisresearch.org/about-the-banded-mongoose-project/

The multiple drivers of specialization and their dynamic quality means that it is important to contextualize specialization. For example, a study on four species of neotropical frogs found varying degrees of specialization across multiple populations of each species [18]. The degree of specialization was dependent on a variety of drivers including predation and both intra- and inter-specific competition. Notably, the direction of the relationship between degree of specialization and each driver was species specific. This study highlights that one species may not always be more specialized than another, but that a populations’ specialization is context dependent.

Therefore, it is important to not only be aware of the degree of specialization present in a population, but to also understand its dynamics and drivers. These relationships can then be used to understand how, and why, a population may react to competition from other species, predators, and changes in resource availability [20].  A population’s specialization can also affect the specialization of other populations and community dynamics [25], therefore, it’s important to consider and study individual specialization on both the population and community level. I am excited to start using our incredible six-year dataset to start investigating these questions for PCFG gray whales, so stay tuned for results!

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References

[1]       M. Taborsky, M. A. Cant, and J. Komdeur, The Evolution of Social Behaviour. Cambridge: Cambridge University Press, 2021. doi: 10.1017/9780511894794.

[2]       E. R. Pianka, “Niche Overlap and Diffuse Competition,” vol. 71, no. 5, pp. 2141–2145, 1974.

[3]       P. Matich et al., “Ecological niche partitioning within a large predator guild in a nutrient-limited estuary,” Limnol. Oceanogr., vol. 62, no. 3, pp. 934–953, 2017, doi: https://doi.org/10.1002/lno.10477.

[4]       S. D. Newsome et al., “The interaction of intraspecific competition and habitat on individual diet specialization: a near range-wide examination of sea otters,” Oecologia, vol. 178, no. 1, pp. 45–59, May 2015, doi: 10.1007/s00442-015-3223-8.

[5]       M. T. Tinker, G. Bentall, and J. A. Estes, “Food limitation leads to behavioral diversification and dietary specialization in sea otters,” Proc. Natl. Acad. Sci., vol. 105, no. 2, pp. 560–565, Jan. 2008, doi: 10.1073/pnas.0709263105.

[6]       M. T. Tinker, M. Mangel, and J. A. Estes, “Learning to be different: acquired skills, social learning, frequency dependence, and environmental variation can cause behaviourally mediated foraging specializations,” Evol. Ecol. Res., vol. 11, pp. 841–869, 2009.

[7]       M. T. Tinker et al., “Structure and mechanism of diet specialisation: testing models of individual variation in resource use with sea otters,” Ecol. Lett., vol. 15, no. 5, pp. 475–483, 2012, doi: 10.1111/j.1461-0248.2012.01760.x.

[8]       S. Rossman et al., “Foraging habits in a generalist predator: Sex and age influence habitat selection and resource use among bottlenose dolphins (Tursiops truncatus),” Mar. Mammal Sci., vol. 31, no. 1, pp. 155–168, 2015, doi: https://doi.org/10.1111/mms.12143.

[9]       L. G. Torres, “A kaleidoscope of mammal , bird and fish : habitat use patterns of top predators and their prey in Florida Bay,” vol. 375, pp. 289–304, 2009, doi: 10.3354/meps07743.

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The costs and benefits of automated behavior classification

Clara Bird, PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

“Why don’t you just automate it?” This is a question I am frequently asked when I tell someone about my work. My thesis involves watching many hours of drone footage of gray whales and meticulously coding behaviors, and there are plenty of days when I have asked myself that very same question. Streamlining my process is certainly appealing and given how wide-spread and effective machine learning methods have become, it is a tempting option to pursue. That said, machine learning is only appropriate for certain research questions and scales, and it’s important to consider these before investing in using a new tool.

The application of machine learning methods to behavioral ecology is called computational ethology (Anderson & Perona, 2014). To identify behaviors from videos, the model tracks individuals across video frames and identifies patterns of movement that form a behavior. This concept is similar to the way we identify a whale as traveling if it’s moving in a straight line and as foraging if it’s swimming in circles within a small area (Mayo & Marx, 1990, check out this blog to learn more). The level of behavioral detail that the model is able to track  depends on the chosen method (Figure 1, Pereira et al., 2020). These methods range from tracking each animal as a simple single point (called a centroid) to tracking the animal’s body positioning in 3D (this method is called pose estimation), which range from providing less detailed to more detailed behavior definitions. For example, tracking an individual as a centroid could be used to classify traveling and foraging behaviors, while pose estimation could identify specific foraging tactics. 

Figure 1. Figure from Pereira et al. (2020) illustrating the different methods of animal behavior tracking that are possible using machine learning.

Pose estimation involves training the machine learning algorithm to track individual anatomical features of an individual (e.g., the head, legs, and tail of a rat), meaning that it can define behaviors in great detail. A behavior state could be defined as a combination of the angle between the tail and the head, and the stride length. 

For example, Mearns et al. (2020) used pose estimation to study how zebrafish larvae in a lab captured their prey. They tracked the tail movements of individual larvae when presented with prey and classified these movements into separate behaviors that allowed them to associate specific behaviors with prey capture (Figure 2). The authors found that these behaviors occurred in a specific sequence, that the behaviors kept the prey within the larvae’s line of sight, and that the sequence was triggered by visual cues.  In fact, when they removed the visual cue of the prey, the larvae terminated the behavior sequence, meaning that the larvae are continually choosing to do each behavior in the sequence, rather than the sequence being one long behavior event that is triggered only by the initial visual cue. This study is a good example of the applicability of machine learning models for questions aimed at kinematics and fine-scale movements. Pose estimation has also been used to study the role of facial expression and body language in rat social communication (Ebbesen & Froemke, 2021). 

Figure 2. Excerpt from figure 1 of Mearns et al. (2020) illustrating (A) the camera set up for their experiment, (B) how the model tracked the eye angles and tail of the larvae fish, (C) the kinematics extracted from the footage. In panel (C) the top plot shows how the eyes converged on the same object (the prey) during prey capture event, the middle plot shows when the tail was curved to the left or the right, and the bottom plot shows the angle of the tail tip relative to the body.

While previous machine learning methods to track animal movements required individuals to be physically marked, the current methods can perform markerless tracking (Pereira et al., 2020). This improvement has broadened the kinds of studies that are possible. For example, Bozek et al., (2021) developed a model that tracked individuals throughout an entire honeybee colony and showed that certain individual behaviors were spatially distributed within the colony (Figure 3). Machine learning enabled the researchers to track over 1000 individual bees over several months, a task that would be infeasible for someone to do by hand. 

Figure 3. Excerpt from figure 1 of Bozek et al., (2021) showing how individual bees and their trajectories were tracked.

These studies highlight that the potential benefits of using machine learning when studying fine scale behaviors (like kinematics) or when tracking large groups of individuals. Furthermore, once it’s trained, the model can process large quantities of data in a standardized way to free up time for the scientists to focus on other tasks.

While machine learning is an exciting and enticing tool, automating behavior detection via machine learning could be its own PhD dissertation. Like most things in life, there are costs and benefits to using this technique. It is a technically difficult tool, and while applications exist to make it more accessible, knowledge of the computer science behind it is necessary to apply it effectively and correctly. Secondly, it can be tedious and time consuming to create a training dataset for the model to “learn” what each behavior looks like, as this step involves manually labeling examples for the model to use. 

As I’ve mentioned in a previous blog, I came quite close to trying to study the kinematics of gray whale foraging behaviors but ultimately decided that counting fluke beats wasn’t necessary to answer my behavioral research questions. It was important to consider the scale of my questions (as described in Allison’s blog) and I think that diving into more fine-scale kinematics questions could be a fascinating follow-up to the questions I’m asking in my PhD. 

For instance, it would be interesting to quantify how gray whales use their flukes for different behavior tactics. Do gray whales in better body condition beat their flukes more frequently while headstanding? Does the size of the fluke affect how efficiently they can perform certain tactics? While these analyses would help quantify the energetic costs of different behaviors in better detail, they aren’t necessary for my broad scale questions. Consequently, taking the time to develop and train a pose estimation machine learning model is not the best use of my time.

That being said, I am interested in applying machine learning methods to a specific subset of my dataset. In social behavior, it is not only useful to quantify the behaviors exhibited by each individual but also the distance between them. For example, the distance between a mom and her calf can be indicative of the calves’ dependence on its mom (Nielsen et al., 2019). However, continuously measuring the distance between two individuals throughout a video is tedious and time intensive, so training a machine learning model could be an effective use of time. I plan to work with an intern this summer to develop a machine learning model to track the distance between pairs of gray whales in our drone footage and then relate this distance data with the manually coded behaviors to examine patterns in social behavior (Figure 4).  Stay tuned to learn more about our progress!

Figure 4. A mom and calf pair surfacing together. Image collected under NOAA/NMFS permit #21678

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References

Anderson, D. J., & Perona, P. (2014). Toward a Science of Computational Ethology. Neuron, 84(1), 18–31. https://doi.org/10.1016/j.neuron.2014.09.005

Bozek, K., Hebert, L., Portugal, Y., Mikheyev, A. S., & Stephens, G. J. (2021). Markerless tracking of an entire honey bee colony. Nature Communications, 12(1), 1733. https://doi.org/10.1038/s41467-021-21769-1

Ebbesen, C. L., & Froemke, R. C. (2021). Body language signals for rodent social communication. Current Opinion in Neurobiology, 68, 91–106. https://doi.org/10.1016/j.conb.2021.01.008

Mayo, C. A., & Marx, M. K. (1990). Surface foraging behaviour of the North Atlantic right whale, Eubalaena glacialis , and associated zooplankton characteristics. Canadian Journal of Zoology, 68(10), 2214–2220. https://doi.org/10.1139/z90-308

Mearns, D. S., Donovan, J. C., Fernandes, A. M., Semmelhack, J. L., & Baier, H. (2020). Deconstructing Hunting Behavior Reveals a Tightly Coupled Stimulus-Response Loop. Current Biology, 30(1), 54-69.e9. https://doi.org/10.1016/j.cub.2019.11.022

Nielsen, M., Sprogis, K., Bejder, L., Madsen, P., & Christiansen, F. (2019). Behavioural development in southern right whale calves. Marine Ecology Progress Series, 629, 219–234. https://doi.org/10.3354/meps13125

Pereira, T. D., Shaevitz, J. W., & Murthy, M. (2020). Quantifying behavior to understand the brain. Nature Neuroscience, 23(12), 1537–1549. https://doi.org/10.1038/s41593-020-00734-z

Of snakes and whales: How food availability and body condition affect reproduction

Clara Bird, PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Over six field seasons the GEMM lab team has conducted nearly 500 drone flights over gray whales, equaling over 100 hours of footage. These hours of footage are the central dataset for my PhD dissertation, so it’s up to me to process them all. This process can be challenging, tedious, and daunting, but it is also quite fun and a privilege to be the one person who gets to watch all the footage. It’s fascinating to get to know the whales and their behaviors and pick up on patterns. It motivates me to get through this video processing step and start doing the data analysis. Recently, it’s been especially fun to notice patterns that I’ve seen mentioned in the literature. One example is adult social behavior. 

There are two categories of social behavior that I’m interested in studying: maternal behavior, defined as interactions between a mom and its calf, and general social behaviors, defined as social interactions between non-mom/calf pairs. In this blog I’ll focus on general social behaviors, but if you’re interested in maternal behavior check out this blog. General social behavior, which I’ll refer to as social behavior moving forward, includes tactile interactions and promiscuous behaviors (Torres et al. 2018; Clip 1). While gray whales in the PCFG range are primarily foraging, researchers have observed increases in social behavior towards the end of the foraging season (Stelle et al., 2008; Torres et al., 2018). We think that this indicates that the whales are starting to focus less on feeding and more on breeding. This tradeoff of foraging vs. socializing time is interesting because it comes at an energetic cost.

Clip 1. Example of social interaction between a male and female gray whale off the coast of Oregon, USA. Collected under NOAA/NMFS permit #21678

Broadly, animals need to balance the energetic demands of survival with those of reproduction. They need to reproduce to pass on their genes, but reproduction is energetically demanding, and animals also need to survive and grow to be able to reproduce. The decision to reproduce is costly because reproduction requires energetic investment and time investment since animals do not forage (gaining energy) when they are socializing. Consequently, only animals with sufficient energy reserves (i.e., body condition) to invest in reproduction actually engage in reproduction. Given these costs associated with reproduction, we expect to see a relationship between social behavior and body condition (Green, 2001) with mainly animals in good body condition engaging in social behavior because these animals have sufficient reserves to sustain the cost. Furthermore, since body condition is an indicator of foraging success and prey availability, environmental conditions can also affect social behavior and reproduction through this pathway. 

Rahman et al. (2014) used a lab experiment to study the relationship between nutritional stress and male guppy courtship behavior (Figure 1). In their experiment they tested for the effects of both decreased diet quantity and quality on the frequency of male courtship behaviors. Rahman et al (2014) found that individuals in the low-quantity group were significantly smaller than those in the high-quality group and that diet quantity had a significant effect on the frequency of courtship behaviors. Males fed a low-quantity diet performed fewer courtship behaviors. Interestingly, there was no significant effect of diet quality on courtships behavior, although there was some evidence of an interaction effect, which suggests that within the low-quantity group, males fed with high-quality food performed more courtship behaviors that those fed with low-quality food. This study is interesting because it shows how foraging success (diet quantity and quality) can affect courting behavior. 

Figure 1. A guppy (Rahman et al., 2013)

However, guppies are not the ideal species for comparison to gray whales because gray whales and guppies have quite different life history traits. A more fitting comparison would be with an example species with more in common with gray whales, such as viviparous capital breeders. Viviparous animals develop the embryo inside the body and give live birth. Capital breeders forage to build energy reserves and then rely on those energy reserves during reproduction. Surprisingly, I found asp vipers to be a good example species for comparison to gray whales.

Asp vipers (Figure 2) are viviparous snakes who are considered capital breeders because they forage prior to hibernation, and then begin reproduction immediately following hibernation without additional foraging. Naulleau & Bonnet (1996) conducted a field study on female asp vipers to determine if there was a difference in body condition at the start of the breeding season between females who reproduced or not during that season. To do this they marked individuals and measured their body condition at the start of the breeding season and then recaptured those individuals at the end of the breeding season and recorded whether the individual had reproduced. Interestingly, they found that there was a strongly significant difference in body condition between females that did and did not reproduce. In fact, they discovered that no female below a certain body condition value reproduced, meaning that they found a body condition threshold for reproduction. 

Figure 2. An asp viper

Additionally, a study on water pythons found that their body condition threshold for reproduction shifted over time in response to prey availability (Madsen & Shine, 1999). These authors found that females lowered their threshold after several consecutive years of poor prey availability. These studies are really exciting to me because they address questions that the GRANITE project team is interested in tackling.

Understanding the relationship between body condition and reproduction in gray whales is an important puzzle piece for our work. The aim of the GRANITE project is to understand how the effects of stressors on individual whales scales up to population level impacts (read Lisa’s blog to learn more). Reproduction rates play a big role in population dynamics, so it is important to understand what factors affect reproduction. Since we’re studying these whales on their foraging grounds, assessing body condition provides an important link between foraging behavior and reproduction. 

For example, if an individual’s response to a stressor is to forage less, that may lead to poorer body condition, meaning that they may be less likely to reproduce. While reduced reproduction in one individual may not have a big effect on the population, the same response from multiple individuals could impact the population’s dynamics (i.e., increasing or decreasing abundance). Understanding these different relationships between behavior, body condition, and reproduction rates is a big undertaking, but it’s exciting to be a member of the GRANITE team as this strong group of scientists works to bring together different data streams to work on this big picture question. We’re all deep into data processing right now so stay tuned over the next few years to learn more about gray whale social behavior and to find out if fat whales are more social than skinny whales. 

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References

Green, A. J. (2001). Mass/Length Residuals: Measures of Body Condition or Generators of Spurious Results? Ecology, 82(5), 1473–1483. https://doi.org/10.1890/0012-9658(2001)082[1473:MLRMOB]2.0.CO;2

Madsen, T., & Shine, R. (1999). The adjustment of reproductive threshold to prey abundance in a capital breeder. Journal of Animal Ecology, 68(3), 571–580. https://doi.org/10.1046/j.1365-2656.1999.00306.x

Naulleau, G., & Bonnet, X. (1996). Body Condition Threshold for Breeding in a Viviparous Snake. Oecologia, 107(3), 301–306.

Rahman, M. M., Kelley, J. L., & Evans, J. P. (2013). Condition-dependent expression of pre- and postcopulatory sexual traits in guppies. Ecology and Evolution, 3(7), 2197–2213. https://doi.org/10.1002/ece3.632

Rahman, M. M., Turchini, G. M., Gasparini, C., Norambuena, F., & Evans, J. P. (2014). The Expression of Pre- and Postcopulatory Sexually Selected Traits Reflects Levels of Dietary Stress in Guppies. PLOS ONE, 9(8), e105856. https://doi.org/10.1371/journal.pone.0105856

Stelle, L. L., Megill, W. M., & Kinzel, M. R. (2008). Activity budget and diving behavior of gray whales (Eschrichtius robustus) in feeding grounds off coastal British Columbia. Marine Mammal Science, 24(3), 462–478. https://doi.org/10.1111/j.1748-7692.2008.00205.x

Torres, L. G., Nieukirk, S. L., Lemos, L., & Chandler, T. E. (2018). Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science, 5(SEP). https://doi.org/10.3389/fmars.2018.00319

Memoirs from above: drone observations of blue, humpback, Antarctic minke, and gray whales

By KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

With the GRANITE field season officially over, we are now processing all of the data we collected this summer. For me, I am starting to go through all the drone videos to take snapshots of each whale to measure their body condition. As I go through these videos, I am reflecting on the different experiences I am fortunate enough to have with flying different drones, in different environments, over different species of baleen whales: blue, humpback, Antarctic minke, and now gray whales. Each of these species have a different morphological design and body shape (Woodward et al., 2006), which leads to different behaviors that are noticeable from the drone. Drones create immense opportunity to learn how whales thrive in their natural environments [see previous blog for a quick history], and below are some of my memories from above. 

I first learned how drones could be used to study the morphology and behavior of large marine mammals during my master’s degree at Duke University, and was inspired by the early works of John Durban (Durban et al., 2015, 2016) Fredrick Christiansen (Christiansen et al., 2016) and Leigh Torres (Torres et al., 2018). I immediately recognized the value and utility of this technology as a new tool to better monitor the health of marine mammals. This revelation led me to pursue a PhD with the Duke University Marine Robotics and Remote Sensing (MaRRS) Lab led by Dr. Dave Johnston where I helped further develop tools and methods for collecting drone-based imagery on a range of species in different habitats. 

When flying drones over whales, there are a lot of moving parts; you’re on a boat that is moving, flying something that is moving, following something that is moving. These moving elements are a lot to think about, so I trained hard, so I did not have to think about each step and flying felt intuitive and natural. I did not grow up playing video games, so reaching this level of comfort with the controls took a lot of practice. I practiced for hours over the course of months before my first field excursion and received some excellent mentorship and training from Julian Dale, the lead engineer in the MaRRS Lab. Working with Julian and the many hours of training helped me establish a solid foundation in my piloting skills and feel confident working in various environments on different species. 

Blue whales offshore of Monterey, California. 

In 2017 and 2018 I was involved in collaborative project with the MaRRS Lab and Goldbogen Lab at Stanford University, where we tagged and flew drones over blue whales offshore of Monterey, California. We traveled about an hour offshore and reliably found groups of blue whales actively feeding. Working offshore typically brought a large swell, which can often make landing the drone back into your field partner’s hands tricky as everything is bobbing up and down with the oscillations of the swell. Fortunately, we worked from a larger research vessel (~56 ft) and quickly learned that landing the drone in the stern helped dampen the effects of bobbing up and down. The blue whales we encountered often dove to a depth of around 200 m for about 20-minute intervals, then come to the surface for only a few minutes. This short surface period provided only a brief window to locate the whale once it surfaced and quickly fly over it to collect the imagery needed before it repeated its dive cycle. We learned to be patient and get a sense of the animal’s dive cycle before launch in order to time our flights so the drone would be in the air a couple of minutes before the whale surfaced. 

Once over the whales, the streamlined body of the blue whales was noticeable, with their small, high aspect ratio flippers and fluke that make them so well adapted for fast swimming in the open ocean (Fig. 1) (Woodward et al., 2006). I also noticed that because these whales are so large (often 21 – 24 m), I often flew at higher altitudes to be able fit them within the field of view of the camera. It was also always shocking to see how small the tagging boat (~8 m) looked when next to Earth’s largest creatures. 

Figure 1. Two blue whales surface after a deep dive offshore of Monterey, Ca. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03)

Antarctic minke whales and humpback whales along the Western Antarctic PeninsulaA lot of the data included in my dissertation came from work along the Western Antarctic Peninsula (WAP), which had a huge range of weather conditions, from warm and sunny days to cold and snowy/foggy/rainy/windy/icy days. A big focus was often trying to keep my hands warm, as it was often easier to fly without gloves in order to better feel the controls. One of the coldest days I remember was late in the season in mid-June (almost winter!) in Wilhemina Bay where ice completely covered the bay in just a couple hours, pushing the whales out into the Gerlache Strait; I suspect this was the last ice-free day of the season. Surprisingly though, the WAP also brought some of the best conditions I have ever flown in. Humpback and Antarctic minke whales are often found deep within the bays along the peninsula, which provided protection from the wind. So, there were times where it would be blowing 40 mph in the Gerlache Strait, but calm and still in the bays, such as Andvord Bay, which allowed for some incredible conditions for flying. Working from small zodiacs (~7 m) allowed us more maneuverability for navigating around or through the ice deep in the bays (Fig. 2) 

Figure 2. Navigating through ice-flows along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Flying over Antarctic minke whale was always rewarding, as they are very sneaky and can quickly disappear under ice flows or in the deep, dark water. Flying over them often felt like a high-speed chase, as their small streamlined bodies makes them incredibly quick and maneuverable, doing barrel rolls, quick banked turns, and swimming under and around ice flows (Fig. 3). There would often be a group between 3-7 individuals and it felt like they were playing tag with each other – or perhaps with me!  

Figure 3. Two Antarctic minke whales swimming together along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Humpbacks displayed a wide range of behaviors along the WAP. Early in the season they continuously fed throughout the entire day, often bubble net feeding in groups typically of 2-5 animals (Fig. 4). For as large as they are, it was truly amazing to see how they use their pectoral fins to perform quick accelerations and high-speed maneuvering for tight synchronized turns to form bubble nets, which corral and trap their krill, their main food source (Fig. 4) (Woodward et al., 2006). Later in the season, humpbacks switched to more resting behavior in the day and mostly fed at night, taking advantage of the diel vertical migration of krill. This behavior meant we often found humpbacks snoozing at the surface after a short dive, as if they were in a food coma. They also seemed to be more curious and playful with each other and with us later in the season (Fig. 5).

We also encountered a lot of mom and calf pairs along the WAP. Moms were noticeably skinny compared to their plump calf in the beginning of the season due to the high energetic cost of lactation (Fig. 6). It is important for moms to regain this lost energy throughout the feeding season and begin to wean their calves. I often saw moms refusing to give milk to their nudging calf and instead led teaching lessons for feeding on their own.

Figure 4. Two humpback whales bubble-net feeding early in the feeding season (December) along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)
Figure 5. A curious humpback whale dives behind our Zodiac along the Western Antarctic Peninsula. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)
Figure 6. A mom and her calf rest at the surface along the Western Antarctic Peninsula. Note how the mom looks skinnier compared to her plump calf, as lactation is the most energetically costly phase of the reproductive cycle. (Image credit: Duke University Marine Robotics and Remote Sensing under NOAA permit 14809-03 and ACA permits 2015-011 and 2020-016.)

Gray whales off Newport, Oregon

All of these past experiences helped me quickly get up to speed and jump into action with the GRANITE field team when I officially joined the GEMM Lab this year in June. I had never flown a DJI Inspire quadcopter before (the drone used by the GEMM Lab), but with my foundation piloting different drones, some excellent guidance from Todd and Clara, and several hours of practice to get comfortable with the new setup, I was flying over my first gray whale by day three of the job. 

The Oregon coast brings all sorts of weather, and some days I strangely found myself wearing a similar number of layers as I did in Antarctica. Fog, wind, and swell could all change within the hour, so I learned to make the most of weather breaks when they came. I was most surprised by how noticeably different gray whales behave compared to the blue, Antarctic minke, and humpback whales I had grown familiar with watching from above. For one, it is absolutely incredible to see how these huge whales use their low-aspect ratio flippers and flukes (Woodward et al., 2006) to perform low-speed, highly dynamic maneuvers to swim in very shallow water (5-10 m) so close to shore (<1m sometimes!) and through kelp forest or surf zones close to the beach. They have amazing proprioception, or the body’s ability to sense its movement, action, and position, as gray whales often use their pectoral fins and fluke to stay in a head standing position (see Clara Bird’s blog) to feed in the bottom sediment layer, all while staying in the same position and resisting the surge of waves that could smash them against the rocks (Video 1) . It is also remarkable how the GEMM Lab knows each individual whale based on natural skin marks, and I started to get a better sense of each whale’s behavior, including where certain individuals typically like to feed, or what their dive cycle might be depending on their feeding behavior. 

Video 1. Two Pacific Coast Feeding Group (PCFG) gray whales “head-standing” in shallow waters off the coast of Newport, Oregon. NOAA/NMFS permit #21678

I feel very fortunate to be a part of the GRANITE field team and to contribute to data collection efforts. I look forward to the data analysis phase to see what we learn about how the morphology and behavior of these gray whales to help them thrive in their environment. 

References: 

Christiansen, F., Dujon, A. M., Sprogis, K. R., Arnould, J. P. Y., and Bejder, L. (2016).Noninvasive unmanned aerial vehicle provides estimates of the energetic cost of reproduction in humpback whales. Ecosphere 7, e01468–18.

Durban, J. W., Fearnbach, H., Barrett-Lennard, L. G., Perryman, W. L., & Leroi, D. J. (2015). Photogrammetry of killer whales using a small hexacopter launched at sea. Journal of Unmanned Vehicle Systems, 3(3), 131-135.

Durban, J. W., Moore, M. J., Chiang, G., Hickmott, L. S., Bocconcelli, A., Howes, G., et al.(2016). Photogrammetry of blue whales with an unmanned hexacopter. Mar. Mammal Sci. 32, 1510–1515.

Torres, L. G., Nieukirk, S. L., Lemos, L., & Chandler, T. E. (2018). Drone up! Quantifying whale behavior from a new perspective improves observational capacity. Frontiers in Marine Science, 5, 319.

Woodward, B. L., Winn, J. P., and Fish, F. E. (2006). Morphological specializations of baleen whales associated with hydrodynamic performance and ecological niche. J. Morphol. 267, 1284–1294.

Learning the right stuff – examining social transmission in humans, monkeys, and cetaceans

Clara Bird, PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

The start of a new school year is always an exciting time. Like high school, it means seeing friends again and the anticipation of preparing to learn something new. Even now, as a grad student less focused on coursework, the start of the academic year involves setting project timelines and goals, most of which include learning. As I’ve been reflecting on these goals, one of my dad’s favorite sayings has been at the forefront of my mind. As an overachieving and perfectionist kid, I often got caught up in the pursuit of perfect grades, so the phrase “just learn the stuff” was my dad’s reminder to focus on what matters. Getting good grades didn’t matter if I wasn’t learning. While my younger self found the phrase rather frustrating, I have come to appreciate and find comfort in it. 

Given that my research is focused on behavioral ecology, I’ve also spent a lot of time thinking about how gray whales learn. Learning is important, but also costly. It involves an investment of energy (a physiological cost, Christie & Schrater, 2015; Jaumann et al., 2013), and an investment of time (an opportunity cost). Understanding the costs and benefits of learning can help inform conservation efforts because how an individual learns today affects the knowledge and tactics that the individual will use in the future. 

Like humans, individual animals can learn a variety of tactics in a variety of ways. In behavioral ecology we classify the different types of learning based on the teacher’s role (even though they may not be consciously teaching). For example, vertical transmission is a calf learning from its mom, and horizontal transmission is an individual learning from other conspecifics (individuals of the same species) (Sargeant & Mann, 2009). An individual must be careful when choosing who to learn from because not all strategies will be equally efficient. So, it stands to reason than an individual should choose to learn from a successful individual. Signals of success can include factors such as size and age. An individual’s parent is an example of success because they were able to reproduce (Barrett et al., 2017). Learning in a population can be studied by assessing which individuals are learning, who they are learning from, and which learned behaviors become the most common.

An example of such a study is Barrett et al. (2017) where researchers conducted an experiment on capuchin monkeys in Costa Rica. This study centered around the Panama ́fruit, which is extremely difficult to open and there are several documented capuchin foraging tactics for processing and consuming the fruit (Figure 1). For this study, the researchers worked with a group of monkeys who lived in a habitat where the fruit was not found, but the group included several older members who had learned Panamá fruit foraging tactics prior to joining this group. During a 75-day experiment, the researchers placed fruits near the group (while they weren’t looking) and then recorded the tactics used to process the fruit and who used each tactic. Their results showed that the most efficient tactic became the most common tactic over time, and that age-bias was a contributing factor, meaning that individuals were more like to copy older members of the group. 

Figure 1. Figure from Barrett et al. (2017) showing a capuchin monkey eating a Panamá fruit using the canine seam technique.

Social learning has also been documented in dolphin societies. A long-term study on wild bottlenose dolphins in Shark Bay, Australia assessed how habitat characteristics and the foraging behaviors used by moms and other conspecifics affected the foraging tactics used by calves (Sargeant & Mann, 2009). Interestingly, although various factors predicted what foraging tactic was used, the dominant factor was vertical transmission where the calf used the tactic learned from its mom (Figure 2). Overall, this study highlights the importance of considering a variety of factors because behavioral diversity and learning are context dependent.

Figure 2. Figure from Sargeant & Mann (2009) showing that the probability of a calf using a tactic was higher if the mother used that tactic.

Social learning is something that I am extremely interested in studying in our study population of gray whales in Oregon. While studies on social learning for such long-lived animals require a longer study period than of the span of our current dataset, I still find it important to consider the role learning may play. One day I would love to delve into the different factors of learning by these gray whales and answer questions such as those addressed in the studies I described above. Which foraging tactics are learned? How much of a factor is vertical transmission? Considering that gray whale calves spend the first few months of the foraging season with their mothers I would expect that there is at least some degree of vertical transmission present. Furthermore, how do environmental conditions affect learning? What tactics are learned in good vs. poor years of prey availability? Does it matter which tactic is learned first? While the chances that I’ll get to address these questions in the next few years are low, I do think that investigating how tactic diversity changes across age groups could be a good place to start. As I’ve discussed in a previous blog, my first dissertation chapter will focus on quantifying the degree of individual specialization present in my study group. After reading about age-biased learning, I am curious to see if older whales, as a group, use fewer tactics and if those tactics are the most energetically efficient.

The importance of understanding learning is related to that of studying individual specialization, which can allows us to estimate how behavioral tactics might change in popularity over time and space. We could then combine this with knowledge of how tactics are related to morphology and habitat and the associated energetic costs of each tactic. This knowledge would allow us to estimate the impacts of environmental change on individuals and the population. While my dissertation research only aims to provide a few puzzle pieces in this very large and complicated gray whale ecology puzzle, I am excited to see what I find. Writing this blog has both inspired new questions and served as a good reminder to be more patient with myself because I am still, “just learning the stuff”.

The early phases of studying harbor seal pup behavior along the Oregon coast

By Miranda Mayhall, Masters Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Recently, when expected to choose a wildlife species for behavioral observation for one of my Oregon State University graduate courses, I immediately chose harbor seals as my focus. Harbor seals (Fig 1) are an abundant species and in proximity to the Hatfield Marine Science Center (HMSC) (Steingass et al., 2019) where I will be spending much of my time this summer, making logistics easy. Studying pinnipeds (marine mammals with a finned foot, seals, walrus, and sea lions) is appealing due to their undeniably cute physique, floppy nature on land, and super agile nature in the water. I am working to iron out my methods for this study, which I hope to work through in this initial phase of my research project.

Figure 1. Harbor seal hauling out to rest on rocks off Oregon Coast near HMSC.

Behaviors:

At times it can appear that the most interesting harbor seal behaviors occur under water, and the haul out time is simply time for resting. During mating season, most adult seal behaviors take place in the water, such as the incredible vocal acoustics displayed by the males to attract the females (Matthews et al., 2018). However, I hypothesize that young pups can capitalize on haul out time by practicing becoming adults (while the adults are taking that time to rest) and therefore I plan to observe their haul out behaviors in their first summer of life. Specifically, I will document seal pup vocal behavior to evaluate how they are learning to use sound. I am beginning this study in late July, which is just after pupping season (Granquist et al., 2016). This should give me the opportunity to find pups along the Oregon coast near HMSC, so I intend to visit several locations where harbor seals are known to frequently haul out. Knowing that field work and animal behavior is unpredictable, there is no telling what behaviors I will observe on a given day, or if I will see seals at all. Some days I could come home with lots of seal data and great photos, and other days I could come home with little to report. This will be my first hurdle combined with my time limit (strictly completing this observation in the next five weeks). I intend to schedule at least eight hours of field observation at haul-out sites over the next two weeks and will adjust my schedule based on my success in data collection at that point.  

Figure 2. Harbor Seals hauling out on rocks not too far from HMSC.

Timing:

Prior knowledge on harbor seal haul-out sites along the Oregon coast is clearly important for this project’s success, but I must also pay close attention to the tide cycles. During low tides, haul out locations are exposed and occupied by seals. When the tide is high, the seals are less likely to haul-out (Patterson et al., 2008). Furthermore, according to a recent study conducted on harbor seals residing on the Oregon coast, these seals spend on average 71% of their time in the water and will haul-out for the remainder of their time (Steingass et al., 2019). Therefore, it is crucial to maximize my observation time of hauled out pups wisely.

Concerning timing, I also need to observe locations and periods without too many tourists who can get near the haul-out site. As I learned recently, when children show up and start throwing rocks into the water near where harbor seals are swimming, the seals will recede from the area and no longer be available for observation. As an experiment, I waited for the noisy crowds with unchecked children to leave and only myself, my trusty sidekick (my daughter), and one quiet photographer were left on the beach. Once that happened, we noticed more and more seal heads popping up out of the water. Then they came closer and closer to the beach, splashing around doing somersaults visibly on the surface of the water. It was quite a show. I will either need to account for the presence of humans when evaluating seal behavior or assess only periods without disturbance. Seal pups are easily disturbed by humans, so I will keep a non-invasive distance while positioning myself to hear the vocals.

Figure 3. Hauled-out adult harbor seal on the Oregon coast near HMSC. 

Data Collection and Analysis Approach:

The aspect of this project I am still working out is how to quantify pup vocalizations and their associated behaviors. As I mentioned, I will go out each week for eight hours and record each time I notice a pup exhibiting vocal behavior. I will categorize and describe the sound produced by the pup, and document any associated behavior of the pup or behavioral responses from nearby adult seals. Prior research has found that harbor seals are much attuned to vocal behavior. Mother harbor seals learn to quickly distinguish their own pup’s call within a few days of their birth (Sauve et al. 2015). I hypothesize that pups themselves can discern and use vocalizations, and I am excited to watch them develop over the course of my field observations.

Figure 4. Seal pup on the far-left rock, watching the adults as they appear to rest.

References

Granquist, S.M., & Hauksson, E. (2016). Seasonal, meteorological, tidal, and diurnal effects on haul-out patterns of harbour seals (Phoca vitulina) in Iceland. Polar Biology, 39 (12), 2347-2359.

Matthews, L.P., Blades, B., Parks, S. (2018). Female harbor seal (Phoca vitulina) behavioral response to playbacks of underwater male acoustic advertisement displays. PeerJ, 6, e4547.

Patterson, J., Acevedo-Gutierrez, A. (2008). Tidal influence on the haul-out behavior of harbor seals (Phoca vitulina) At all time levels. Northwestern Naturalist, 89 (1), 17-23.

Sauve, C., Beauplet, G., Hammil, M., Charrier, I. (2015). Mother-pup vocal recognition in harbour seals: influence of maternal behavior, pup voice and habitat sound properties. Animal Behavior, 105 (July 2015), 109-120

Steingass, S., Horning, M., Bishop, A. (2019). Space use of Pacific harbor seals (Phoca vitulina richardii) from two haulout locations along the Oregon coast. PloS one. 14 (7), e0219484.

The right tool for the job: examining the links between animal behavior, morphology and habitat

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

In order to understand a species’ distribution, spatial ecologists assess which habitat characteristics are most often associated with a species’ presence. Incorporating behavior data can improve this analysis by revealing the functional use of each habitat type, which can help scientists and managers assign relative value to different habitat types. For example, habitat used for foraging is often more important than habitat that a species just travels through. Further complexity is added when we consider that some species, such as gray whales, employ a variety of foraging tactics on a variety of prey types that are associated with different habitats. If individual foraging tactic specialization is present, different foraging habitats could be valuable to specific subgroups that use each tactic. Consequently, for a population that uses a variety of foraging tactics, it’s important to study the associations between tactics and habitat characteristics.

Lukoschek and McCormick’s (2001) study investigating the spatial distribution of a benthic fish species’ foraging behavior is a great example of combining data on behavior, habitat, and morphology.  They collected data on the diet composition of individual fish categorized into different size classes (small, medium, and large) and what foraging tactics were used in which reef zones and habitat types. The foraging tactics ranged from feeding in the water column to digging (at a range of depths) in the benthic substrate. The results showed that an interesting combination of fish behavior and morphology explained the observed diet composition and spatial distribution patterns. Small fish foraged in shallower water, on smaller prey, and primarily employed the water column and shallow digging tactics. In contrast, large fish foraged in deep water, on larger prey, and primarily fed by digging deeper into the seafloor (Figure 1). This pattern is explained by both morphology and behavior. Morphologically, the size of the feeding apparatus (mouth gape size) affects the size of the prey that a fish can feed on. The gape of the small fish is not large enough to eat the larger prey that large fish are able to consume. Behaviorally, predation risk also affects habitat selection and tactic use. Small fish are at higher risk of being predated on, so they remain in shallow areas where they are more protected from predators and they don’t dig as deep to forage because they need to be able to keep an eye out for predators. Interestingly, while they found a relationship between the morphology of the fish and habitat use, they did not find an association between specific feeding tactics and habitat types.

Figure 1. Figure from Lukoschek and McCormick (2001) showing that small fish (black bar) were found in shallow habitat while large fish (white bar) were found in deep habitat.

Conversely, Torres and Read (2009) did find associations between theforaging tactics of bottlenose dolphins in Florida Bay, FL and habitat type. Dolphins in this bay employ three foraging tactics: herd and chase, mud ring feeding, and deep diving. Observations of the foraging tactics were linked to habitat characteristics and individual dolphins. The study found that these tactics are spatially structured by depth (Figure 2), with deep diving occurring in deep water whereas mud ring feeding occurrs in shallower water. They also found evidence of individual specialization! Individuals that were observed deep diving were not observed mud ring feeding and vice-versa. Furthermore, they found that individuals were found in the habitat type associated with their preferred tactic regardless of whether they were foraging or not. This result indicates that individual dolphins in this bay have a foraging tactic they prefer and tend to stay in the corresponding habitat type. These findings are really intriguing and raise interesting questions regarding how these tactics and specializations are developed or learned. These are questions that I am also interested in asking as part of my thesis.

Figure 2. Figure from Torres and Read (2009) showing that deep diving is associated with deeper habitat while mud ring feeding is associated with shallow habitat.

Both of these studies are cool examples that, combined, exemplify questions I am interested in examining using our study population of Pacific Coast Feeding Group (PCFG) gray whales. Like both studies, I am interested in assessing how specific foraging tactics are associated with habitat types. Our hypothesis is that different prey types live in different habitat types, so each tactic corresponds to the best way to feed on that prey type in that habitat. While predation risk doesn’t have as much of an effect on foraging gray whales as it does on small benthic fish, I do wonder how disturbance from boats could similarly affect tactic preference and spatial distribution. I am also curious to see if depth has an effect on tactic choice by using the morphology data from our drone-based photogrammetry. Given that these whales forage in water that is sometimes as deep as they are long, it stands to reason that maneuverability would affect tactic use. As described in a previous blog, I’m also looking for evidence of individual specialization. It will be fascinating to see how foraging preference relates to space use, habitat preference, and morphology.

These studies demonstrate the complexity involved in studying a population’s relationship to its habitat. Such research involves considering the morphology and physiology of the animals, their social, individual, foraging, and predator-prey behaviors, and the relationship between their prey and the habitat. It’s a bit daunting but mostly really exciting because better understanding each puzzle piece improves our ability to estimate how these animals will react to changing environmental conditions.

While I don’t have any answers to these questions yet, I will be working with a National Science Foundation Research Experience for Undergraduates intern this summer to develop a habitat map of our study area that will be used in this analysis and potentially answer some preliminary questions about PCFG gray whale habitat use patterns. So, stay tuned to hear more about our work this summer!

References

Lukoschek, V., & McCormick, M. (2001). Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): Relationship between foraging behaviour, habitat use, jaw size, and prey selection. Marine Biology, 138(6), 1099–1113. https://doi.org/10.1007/s002270000530

Torres, L. G., & Read, A. J. (2009). 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, 25(4), 797–815. https://doi.org/10.1111/j.1748-7692.2009.00297.x