“What if I’m wrong? What if I make a mistake?” When I began my career after completing my undergraduate degree, these questions echoed constantly in my head as the stakes were raised and my work was taken more seriously. Of course, this anxiety was not new. As a student, my worst fear had been poor performance in class. Post-undergrad, I was facing the possibility of making a mistake that could impact larger research projects and publications.
Gaining greater responsibility and consequences is a fact of life and an intrinsic part of growing up. As I wrap up my third year of graduate school, I’ve been reflecting on how learning to take on this responsibility as a scientist has been a crucial part of my journey thus far.
A scientist’s job is to ask, and try to answer, questions that no one knows the answer to – which is both terrifying and exciting. It feels a bit like realizing that grown-ups don’t have all the answers as a kid. Becoming comfortable with the fact that my work often involves making decisions that no one definitively can say are wrong or right has been one of my biggest challenges of grad school. The important thing to remember, I’ve learned, is that I’m not making wild guesses – I’m being trained to make the best, most informed decisions possible. And, hopefully, with more experience will come greater confidence.
Through grad school I have learned to take on this responsibility both in the field and the lab, although each brings different experiences. In the field, the stakes can feel higher because the decisions we make affect not just the quality of the data, but the safety of the team (which is always the top priority). I felt this most acutely throughout my first summer as a drone pilot. As a pilot, I am responsible for the safety of the team, the drone, and the quality of the data. As a new pilot, I intensely felt this pressure and would come home feeling more exhausted than usual. Now, in my second field season in this role, I’ve become more comfortable and am slowly building confidence in my abilities as I gain more and more experience.
Video 1 – Two gray whales foraging together off Newport, Oregon, USA. I recorded this footage during my first season as a pilot – a flight I’ll never forget! NOAA/NMFS permit #21678.
I have also had a similar experience in the lab. Once it’s time to work on the analysis of a project, I choose how to clean, analyze, and interpret the data. As a young scientist, every step of the process involves learning new skills and making decisions that I don’t feel entirely qualified to make. When I started analysis for my first PhD chapter, I felt overwhelmed by deciding how to standardize my data, what kind of analysis to perform, and what indices to calculate. And, since it’s my first chapter, I felt further overwhelmed by the worry that any decision I made would become a later regret in a future part of my PhD.
Recently, the most daunting decision has been how to standardize my data. For my first chapter, I am investigating individual specialization of gray whale foraging behavior. The results of this question are not only important for conservation, but for my subsequent work (check out these previous blogs from January 2021and April 2022 for more on this research question). While there is a wealth of literature to draw analysis inspiration from, most of these studies use discrete prey capture data, while I am working with continuous behavior data. So, to make my data points comparable to one another, I need to standardize the behavior observation time of each drone flight to account for the potential bias introduced by recording one individual for more time than another. After experiencing an internal roller coaster of having an idea, thinking it through, deciding it was terrible and restarting the cycle, I was reminded that turning to lab mates and collaborators is the best way to work through a problem.
Image 1 – Comic from phdcomics.com, source: https://phdcomics.com/comics/archive.php?comicid=2008
So, I had as many conversations as I could with my advisor, committee members, and peers. My thinking clarified with every conversation, and I gained confidence in the justification behind my decision. I cannot fully express the comfort that comes from hearing a trusted advisor say, “that makes ecological sense to me”. These conversations have also helped me remember that I am not alone in my worry and that I am not failing because I have these doubts. While I may never be 100% convinced that I’ve made the right decision, I feel much better knowing that I’ve talked it through with the brilliant group of scientists around me. And as I enter an analysis-intensive phase of my PhD, I am extremely grateful to have this community around to challenge, advise, and support me.
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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
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“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
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
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”.
Allison Dawn, new GEMM Lab Master’s student, OSU Department of Fisheries, Wildlife and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab
While standing at the Stone Shelter at the Saint Perpetua Overlook in 2016, I took in the beauty of one of the many scenic gems along the Pacific Coast Highway. Despite being an East Coast native, I felt an unmistakable draw to Oregon. Everything I saw during that morning’s hike, from the misty fog that enshrouded evergreens and the ocean with mystery, to the giant banana slugs, felt at once foreign and a place I could call home. Out of all the places I visited along that Pacific Coast road trip, Oregon left the biggest impression on me.
Figure 1. View from the Stone Shelter at the Cape Perpetua Overlook, Yachats, OR. June 2016.
For my undergraduate thesis, which I recently defended in May 2021, I researched blue whale surface interval behavior. Surface interval events for oxygen replenishment and rest are a vital part of baleen whale feeding ecology, as it provides a recovery period before they perform their next foraging dive (Hazen et al., 2015; Roos et al., 2016). Despite spending so much time studying the importance of resting periods for mammals, that 2016 road trip was my last true extended resting period/vacation until, several years later in 2021, I took another road trip. This time it was across the country to move to the place that had enraptured me.
Now that I am settled in Corvallis, I have reflected on my journey to grad school and my recent road trip; both prepared me for a challenging and exciting new chapter as an incoming MSc student within the Marine Mammal Institute (MMI).
Part 1: Journey to Grad School
When I took that photo at the Cape Perpetua Overlook in 2016, I had just finished the first two semesters of my undergraduate degree at UNC Chapel Hill. As a first-generation, non-traditional student those were intense semesters as I made the transition from a working professional to full-time undergrad.
By the end of my freshman year I was debating exactly what to declare as my major, when one of my marine science TA’s, Colleen, (who is now Dr. Bove!), advised that I “collect experiences, not degrees.” I wrote this advice down in my day planner and have never forgotten it. Of course, obtaining a degree is important, but it is the experiences you have that help lead you in the right direction.
That advice was one of the many reasons I decided to participate in the Morehead City Field Site program, where UNC undergraduates spend a semester at the coast, living on the Duke Marine Lab’s campus in Beaufort, NC. During that semester, students take classes to fulfill a marine science minor while participating in hands-on research, including an honors thesis project. The experience of designing, carrying out, and defending my own project affirmed that graduate school in the marine sciences was right for me. As I move into my first graduate TA position this fall, I hope to pay forward that encouragement to other undergraduates who are making decisions about their own future path.
Figure 2. Final slide from my honors thesis defense. UNC undergraduates, and now fellow alumni, who participated in the Morehead City Field Site program in Fall 2018.
Part 2: Taking a Breather
Like the GEMM Lab’s other new master’s student Miranda, my road trip covered approximately 2,900 miles. I was solo for much of the drive, which meant there was no one to argue when I decided to binge listen to podcasts. My new favorite is How To Save A Planet, hosted by marine biologist Dr. Ayana Elizabeth Johnson and Alex Blumberg. At the end of each episode they provide a call to action & resources for listeners – I highly recommend this show to anyone interested in what you can do right now about climate change.
Along my trip I took a stop in Utah to visit my parents. I had never been to a desert basin before and engaged in many desert-related activities: visiting Zion National Park, hiking in 116-degree heat, and facing my fear of heights via cliff jumping.
Figure 3. Sandstone Rocks at Sand Hollow National Park, Hurricane, Utah. June 2021.
My parents wanted to help me settle into my new home, as parents do, so we drove the rest of the way to Oregon together. As this would be their first visit to the state, we strategically planned a trip to Crater Lake as our final scenic stop before heading into Corvallis.
Figure 4. Wizard Island in Crater Lake National Park, Klamath County, OR. June 2021.
This time off was filled with adventure, yet was restorative, and reminded me the importance of taking a break. I feel ready and refreshed for an intense summer of field work.
Part 3: Rested and Ready
Despite accumulating skills to do research in the field over the years, I have yet to do marine mammal field work (or even see a whale in person for that matter.) My mammal research experience included analyzing drone imagery, behind a computer, that had already been captured. As you can imagine, I am extremely excited to join the Port Orford team as part of the TOPAZ/JASPER projects this summer, collecting ecological data on gray whales and their prey. I will be learning the ropes from Lisa Hildebrand and soaking up as much information as possible as I will be taking over as lead for this project next year.
It will take some time before my master’s thesis is fully developed, but it will likely focus on assessing the environmental factors that influence gray whale zooplankton prey availability, and the subsequent impacts on whale movements and health. For five years, the Port Orford project has conducted GoPro drops at 12 sampling stations to collect data on zooplankton relative abundance.
Figures 5 & 6. GEMM GoPro drop stick assembly and footage demonstrating mysid data collection. July 2021.
Paired with this GoPro is a Time-Depth Recorder (TDR) that provides temperature and depth data. The 2021 addition to this GoPro system is a new dissolved oxygen (DO) sensor the GEMM Lab has just acquired. This new piece of equipment will add to the set of parameters we can analyze to describe what and how oceanographic factors drive prey variability and gray whale presence in our study site.My first task as a GEMM Lab student is to get to know this DO sensor, figure out how it works, set it up, test it, attach it to the GoPro device, and prepare it for data collection during the upcoming Port Orford project starting in 1 week!
Figure 7. The GEMM lab’s new RBR solo3 getting ready for Port Orford. July 2021.
Dissolved oxygen plays a vital role in the ocean; however, climate change and increased nutrient loading has caused the ocean to undergo deoxygenation. According to the IUCN’s 2019 Issues Brief, these factors have resulted in an oxygen decline of 2% since the middle of the 20th century, with most of this loss occurring within the first 1000 meters of the ocean. Two percent may not seem like much, but many species have a narrow oxygen threshold and, like pH changes in coral reef systems, even slight changes in DO can have an impact. Additionally, the first 1000 meters of the ocean contains the greatest amount of species richness and biodiversity.
Previous research done in a variety of systems (i.e., estuarine, marine, and freshwater lakes) shows that dissolved oxygen concentrations can have an impact on predator-prey interactions, where low dissolved oxygen results in decreased predation (Abrahams et al., 2007; Breitburg et al., 1997; Domenici et al., 2007; Kramer et al., 1987); and changes in DO also change prey vertical distributions (Decker et al., 2004). In Port Orford, we are interested in understanding the interplay of factors driving zooplankton community distribution and abundance while investigating the trophic interaction between gray whales and their prey.
I have spent some time with our new DO sensor and am looking forward to its first deployments in Port Orford! Stay tuned for updates from the field!
References
Abrahams, M. V., Mangel, M., & Hedges, K. (2007). Predator–prey interactions and changing environments: who benefits?. Philosophical Transactions of the Royal Society B: Biological Sciences, 362(1487), 2095-2104.
Breitburg, D. L., Loher, T., Pacey, C. A., & Gerstein, A. (1997). Varying effects of low dissolved oxygen on trophic interactions in an estuarine food web. Ecological Monographs, 67(4), 489-507.
Decker, M. B., Breitburg, D. L., & Purcell, J. E. (2004). Effects of low dissolved oxygen on zooplankton predation by the ctenophore Mnemiopsis leidyi. Marine Ecology Progress Series, 280, 163-172.
Domenici, P., Claireaux, G., & McKenzie, D. J. (2007). Environmental constraints upon locomotion and predator–prey interactions in aquatic organisms: an introduction.
Hazen, E. L., Friedlaender, A. S., & Goldbogen, J. A. (2015). Blue whales (Balaenoptera musculus) optimize foraging efficiency by balancing oxygen use and energy gain as a function of prey density. Science Advances, 1(9), e1500469.
Kramer, D. L. (1987). Dissolved oxygen and fish behavior. Environmental biology of fishes, 18(2), 81-92.
Roos, M. M., Wu, G. M., & Miller, P. J. (2016). The significance of respiration timing in the energetics estimates of free-ranging killer whales (Orcinus orca). Journal of Experimental Biology, 219(13), 2066-2077.
By Marc Donnelly, Carleton College, GEMM Lab REU Intern
My name is Marc Donnelly (He/Him) and this summer I have the pleasure of working with Clara Bird and Dr. Leigh Torres on a project, within the GRANITE project, that maps the habitat use of Pacific Coast Feeding Group (PCFG) gray whales. This summer, as a National Science Foundation (NSF) Research Experience for Undergraduates (REU) intern, I have the opportunity to learn about the scientific process in action, build relationships with researchers, and pursue my own research project. I am relatively new to the world of research and even more green in the field of marine science. In September, I will start my fourth and last year as an Environmental Studies major at Carleton College in Minnesota, but for the next seven weeks my bread and butter is whales. I could not be more excited about the road ahead. I have read about gray whales, examined pictures of them, and even studied videos of the oblong beauties (Video 1). But the one thing I have not done, and probably will not do this summer, is see one… or a boat for that matter, or a single piece of field equipment. Not in person anyway. This is because I am a remote intern. And before I continue, let me clarify that I am not lamenting unfortunate circumstances. These past three weeks have already been enjoyable, fulfilling and exciting and I expect the summer will only get better. Yet, like with so many people in the past 16 months, my expected role has undergone some changes, so I want to take this opportunity to share my experience so far.
Video 1. Examples of gray whale behaviors observed from the drone
Especially when thinking about engaging with hands-on work, a fundamental aspect of any research program is place. REUs are competitive and sought after positions because they supply undergrads, who have to balance coursework with the desire to fully immerse themselves in a research community, with the opportunity to pursue a genuine research experience. Just being in a room surrounded by peers, grad students, and seasoned scientists who are all bubbling over with excitement and ideas is a fundamentally different (and might I add more motivating) experience than classroom lectures. Location is enough by itself to facilitate the connection between a burgeoning scientist and their research community as well as their work itself. Conducting hands-on fieldwork is also a common, highly sought-after aspect of an REU. However, visiting study sites, collecting data, and experiencing your study organism first hand, are all activities that become impossible when working remotely. So what do you do when you lack the benefits of location?
Well, if you had any sense, you would start by furnishing your apartment, finding a mattress that you can sleep on and a table and a chair that you can work at. But if you are like me, 21 and excited to be living away from home in a new city, you might be so overcome with the idea of adventure that you forget sleeping is important. Beyond furniture, the course of my summer has primarily been in the hands of my new mentors at Oregon State University (OSU), the institution funding my position in the GEMM lab, and thankfully they had a much more robust plan for my summer than I did. Data analysis and an in depth literature review have filled the void where my marine mammal companions could have been. This situation does not mean that analyzing data and diving into the literature are not part of in-person internships as well, or that I am not able to build any sort of connection with the gray whales I study, but my computer screen has certainly taken a more central role in my work. This summer, Geographic Information Systems (GIS) software is my weapon of choice. My goal is to create habitat maps of the coastal waters off Newport where gray whales feed that includes characteristics ranging from the type of surface on the bottom of the ocean (i.e., sand, reef, rock, etc) to more ephemeral features like kelp and prey density. This list of features I will map is dynamic based on the purpose and time scale of the map (month, year, static); so suffice it to say I will be making a lot of maps this summer. Once I have produced these habitat maps, the team and I can compare them with our whale sightings to better understand if and how gray whales use certain areas. This work will help us develop a baseline for gray whale ecology, which will ultimately be used to inform disturbance models and conservation efforts.
Figure 1: Me finally getting a table and chair for work.
After finding a way to move work online, the next step is to somehow engineer a social environment that provides people with a sense of community. As explained to the interns during our first professional development workshop, forming these connections are not just important for combating feelings of isolation, but they may also serve as fruitful professional relationships in the near or distant future. After three terms of online classes and vain attempts at forming meaningful connections via awkward breakout rooms and forced group projects, I was preemptively lowering my expectations for how this summer might unfold. I should not have worried; both the GEMM lab and the greater REU cohort have been extraordinary. It has been such a privilege and joy to meet so many compassionate and involved people. Every week there are numerous opportunities for interns to engage with various groups associated with OSU. From one-on-one meetings with my mentors to laid back “coffee breaks” with folks from Oregon Sea Grant, engrossing interactions abound. I even had the chance to attend the Marine Mammal Institute Monthly Meeting, or MMIMM, which for a newcomer to the world of marine science is both a fascinating and intimidating thing to watch unfold.
One of my favorite virtual gatherings of all was our monthly GEMM lab meeting, this month our activity involved brief presentations introducing ourselves and our research. If you are a fan of this blog and have had the opportunity to explore the happenings of the GEMM lab through this page, then you probably have some context to understand the excitement and curiosity I felt while listening about the current GEMM projects through my zoom screen. I was simultaneously humbled and comforted by the impressiveness of the work being undertaken by this group of researchers. Even though I was just being exposed to five-minute overviews of people’s work, it was daunting to compare my own limited knowledge to that of the other people on the call. Most of them have been studying marine megafauna for five years or more and their passion coupled with their grasp on their work was remarkable. I was also comforted by the descriptions of all these wonderful and intriguing projects because it gave me a sense of achievement. This feeling may sound silly, but just by virtue of being on a zoom call with such passionate scientists I felt relieved. Relieved because it seemed as though this community is what I had been working towards for the last few years. Not necessarily the GEMM lab in particular, but a community of inspiring people who care about each other, their work, and improving the world.
Despite the fact that they do not know what I look like from the shoulders down, my GEMM lab cohort has welcomed me into their midst and provided me with the tools and environment I need to connect and learn. I am grateful.
Clara Bird, PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab
When I thought about what doing fieldwork would be like, before having done it myself, I imagined that it would be a challenging, but rewarding and fun experience (which it is). However, I underestimated both ends of the spectrum. I simultaneously did not expect just how hard it would be and could not imagine the thrill of working so close to whales in a beautiful place. One part that I really did not consider was the pre-season phase. Before we actually get out on the boats, we spend months preparing for the work. This prep work involves buying gear, revising and developing protocols, hiring new people, equipment maintenance and testing, and training new skills. Regardless of how many successful seasons came before a project, there are always new tasks and challenges in the preparation phase.
For example, as the GEMM Lab GRANITE project team geared up for its seventh field season, we had a few new components to prepare for. Just to remind you, the GRANITE (Gray whale Response to Ambient Noise Informed by Technology and Ecology) project’s field season typically takes place from June to mid-October of each year. Throughout this time period the field team goes out on a small RHIB (rigid hull inflatable boat), whenever the weather is good enough, to collect photo-ID data, fecal samples, and drone imagery of the Pacific Coast Feeding Group (PCFG) gray whales foraging near Newport, OR, USA. We use the data to assess the health, ecology and population dynamics of these whales, with our ultimate goal being to understand the effect of ambient noise on the population. As previous blogs have described, a typical field day involves long hours on the water looking for whales and collecting data. This year, one of our exciting new updates is that we are going out on two boats for the first part of the field season and starting our season 10 days early (our first day was May 20th). These updates are happening because a National Science Foundation funded seismic survey is being conducted within our study area starting in June. The aim of this survey is to assess geophysical structures but provides us with an opportunity to assess the effect of seismic noise on our study group by collecting data before, during, and after the survey. So, we started our season early in order to capture the “before seismic survey” data and we are using a two-boat approach to maximize our data collection ability.
While this is a cool opportunistic project, implementing the two-boat approach came with a new set of challenges. We had to find a second boat to use, buy a new set of gear for the second boat, figure out the best way to set up our gear on a boat we had not used before, and update our data processing protocols to include data collected from two boats on the same day. Using two boats also means that everyone on the core field team works every day. This core team includes Leigh (lab director/fearless leader), Todd (research assistant), Lisa (PhD student), Ale (new post-doc), and me (Clara, PhD student). Leigh and Todd are our experts in boat driving and working with whales, Todd is our experienced drone pilot, I am our newly certified drone pilot, and Lisa, Ale, and myself are boat drivers. Something I am particularly excited about this season is that Lisa, Ale, and I all have at least one field season under our belts, which means that we get to become more involved in the process. We are learning how to trailer and drive the boats, fly the drones, and handling more of the post-field work data processing. We are becoming more involved in every step of a field day from start to finish, and while it means taking on more responsibility, it feels really exciting. Throughout most of graduate school, we grow as researchers as we develop our analytical and writing skills. But it’s just as valuable to build our skillset for field work. The ocean conditions were not ideal on the first day of the field season, so we spent our first day practicing our field skills.
For our “dry run” of a field day, we went through the process of a typical day, which mostly involved a lot of learning from Leigh and Todd. Lisa practiced her trailering and launching of the boat (figure 1), Ale and Lisa practiced driving the boat, and I practiced flying the drone (figure 2). Even though we never left the bay or saw any whales, I thoroughly enjoyed our dry run. It was useful to run through our routine, without rushing, to get all the kinks out, and it also felt wonderful to be learning in a supportive environment. Practicing new skills is stressful to say the least, especially when there is expensive equipment involved, and no one wants to mess up when they’re being watched. But our group was full of support and appreciation for the challenges of learning. We cheered for successful boat launchings and dockings, and drone landings. I left that day feeling good about practicing and improving my drone piloting skills, full of gratitude for our team and excited for the season ahead.
Figure 1. Lisa (driving the truck) launching the boat.
Figure 2. Clara (seated, wearing a black jacket) landing the drone in Ale’s hands.
All the diligent prep work paid off on Saturday with a great first day (figure 3). We conducted five GoPro drops (figure 4), collected seven fecal samples from four different whales (figure 5), and flew four drone flights over three individuals including our star from last season, Sole. Combined, we collected two trifectas (photo-ID images, fecal samples, and drone footage)! Our goal is to get as many trifectas as possible because we use them to study the relationship between the drone data (body condition and behavior) and the fecal sample data (hormones). We were all exhausted after 10 hours on the water, but we were all very excited to kick-start our field season with a great day.
Figure 3. Lisa on the bow pulpit during our first sighting of the day.
Figure 4. Lisa doing a GoPro drop, she’s lowering the GoPro into the water using the line in her hands.
Figure 5. Clara and Ale collecting a fecal sample.
On Sunday, just one boat went out to collect more data from Sole after a rainy morning and I successfully flew over her from launching to landing! We have a long season ahead, but I am excited to learn and see what data we collect. Stay tuned for more updates from team GRANITE as our season progresses!
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
Clara Bird, PhD Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab
When I started working on my thesis, I anticipated many challenges related to studying the behavioral ecology of gray whales. From processing five-plus years of drone footage to data analysis, there has been no shortage of anticipated and unexpected issues. I recently hit an unexpected challenge when I started video processing that piqued my interest. As I’ve discussed in a previous blog, ethograms are lists of defined behaviors that help us properly and consistently collect data in a standardized approach. Ethograms form a crucial foundation of any behavior study as the behaviors defined ultimately affect what questions can be asked and what patterns are detected. Since I am working off of the thorough ethogram of Oregon gray whales from Torres et al. (2018), I had not given much thought to the process of adding behaviors to the ethogram. But, while processing the first chunk of drone videos, I noticed some behaviors that were not in the original ethogram and struggled to decide whether or not to add them. I learned that ethogram development can lead down several rabbit holes. The instinct to try and identify every movement is strong but dangerous. Every minute movement does not necessarily need to be included and it’s important to remember the ultimate goal of the analysis to avoid getting bogged down.
Fundamental behavior questions cannot be answered without ethograms. For example, Baker et al. (2017) developed an ethogram for bottlenose dolphins in Ireland in order to conduct an initial quantitative behavior analysis. They did so by reviewing published ethograms for bottlenose dolphins, consulting with multiple experts, and revising the ethogram throughout the study. They then used their data to test inter-observer variability, calculate activity budgets, and analyze how the activity budgets varied across space and time.
Howe et al. (2015) also developed an ethogram in order to conduct quantitative behavior analyses. Their goals were to use the ethogram and subsequent analyses to better understand the behavior of beluga whales in Cook Inlet, AK, USA and to inform conservation. They started by writing down all behaviors they observed in the field, then they consolidated their notes into a formal ethogram that they used and refined during subsequent field seasons. They used their data to analyze how the frequencies of different behaviors varied throughout the study area at different times. This study served as an initial analysis investigating the effect of anthropogenic disturbance and was refined in future studies.
My research is similarly geared towards understanding behavior patterns to ultimately inform conservation. The primary questions of my thesis involve individual specialization, patterns of behavior across space, the relationship between behavior and body condition, and social behavior (check out this blog to learn more). While deciding what behaviors to add to my ethogram I’ve had to remind myself of these main questions and the bigger picture. The drone footage lets us see so much detail that it’s tempting to try to define every movement we can observe. One rabbit hole I’ve had to avoid a few times is locomotion. From the footage, it is possible to document fluke beats and pectoral fin strokes. While it could be interesting to investigate how different whales move in different ways, it could easily become a complicated mess of classifying different movements and take me deep into the world of whale locomotion. Talking through what that work would look like reminded me that we cannot answer every question and trying to assess all exciting side projects can cause us to lose focus on the main questions.
While I avoided going down the locomotion rabbit hole, there were some new behaviors that I did add to my ethogram. I’ll illustrate the process with the examples of two new behaviors I recently added: fluke swish and pass under (Clips 1 and 2). Clip 1 shows a whale rapidly moving its fluke to the side. I chose to add fluke swish because it’s such a distinct movement and I’m curious to see if there’s a pattern across space, time, individual, or nearby human activity that might explain its function. Clip 2 shows a calf passing under its mom. It’s not nursing because the calf doesn’t spend time under its mom, it just crosses underneath her. The calf pass under behavior could be a type of mom-calf tactile interaction. Analyzing how the frequency of this behavior changes over time could show how a calf’s dependency on its mom changes over as it ages.
In defining these behaviors, I had to consider how many different variations of this behavior would be included in the definition. This process involves considering at what point a variation of that behavior could serve a different function, even without knowing the function of the original behavior. For fluke swish this process involved deciding to only count a behavior as a fluke swish if it was a big, fast movement. A small and slow movement of the fluke a little to the side could serve a different function, such as turning, or be a random movement.
Clip 1: Fluke swish behavior (Video filmed under NOAA/NMFS research permit #16111 by certified drone pilot Todd Chandler).
Clip 2: Pass under behavior (Video filmed under NOAA/NMFS research permit #16111 by certified drone pilot Todd Chandler).
The next step involved deciding if the behavior would be a ‘state’ or ‘point’ event. A state event is a behavior with a start and stop moment; a point event is instantaneous and assigned to just a point in time. I would categorize a behavior as a state event if I was interested in questions about its duration. For example, I could ask “what percentage of the total observation time was spent in a certain behavior state?” A point event would be a behavior where duration is not applicable, but I could ask a question like “Did whale 1 perform more point event A than whale 2?”. Both fluke swish and pass under are point events because they only happen for an instant. In a pass under the calf is passing under its mom for just a brief point in time, making it a point event. The final step was to name the behavior. As I discussed in this blog, the name of the behavior does not matter as much as the definition but it is important that the name is clear and descriptive. We chose the name fluke swish because the fluke rapidly moves from side to side and pass under because the calf crosses under its mom.
Frankly, in the beginning, I was a bit overwhelmed by the realization that the content of my ethogram would ultimately control the questions I could answer. I could not help but worry that after processing all the videos, I would end up regretting not defining more behaviors. However, after reading some of the literature, chatting with Leigh, and reviewing the initial chunk of videos several times, I am more confidence in my judgment and my ethogram. I have accepted the fact that I can’t anticipate everything, and I am confident that the behaviors I need to answer my research questions are included. The process of reviewing and updating my ethogram has been a rewarding challenge that resulted in a valuable lesson that I will take with me for the rest of my career.
References
Baker, I., O’Brien, J., McHugh, K., & Berrow, S. (2017). An ethogram for bottlenose dolphins (Tursiops truncatus) in the Shannon Estuary, Ireland. Aquatic Mammals, 43(6), 594–613. https://doi.org/10.1578/AM.43.6.2017.594
Howe, M., Castellote, M., Garner, C., McKee, P., Small, R. J., & Hobbs, R. (2015). Beluga, Delphinapterus leucas, ethogram: A tool for cook inlet beluga conservation? Marine Fisheries Review, 77(1), 32–40. https://doi.org/10.7755/MFR.77.1.3
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
