An ‘X’travaganza! Introducing the Marine Mammal Institute’s Center of Drone Excellence (CODEX)

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

Drones are becoming more and more prevalent in marine mammal research, particularly for non-invasively obtaining morphological measurements of cetaceans via photogrammetry to identify important health metrics (see this and this previous blog). For example, the GEMM Lab uses drones for the GRANITE Project to study Pacific Coast Feeding Group (PCFG) gray whales and we have found that PCFG whales are skinnier and morphologically shorter with smaller skulls and flukes compared to the larger Eastern North Pacific (ENP) population. The GEMM Lab has also used drones to document variation in body condition across years and within a season, to diagnose pregnancy, and even measure blowholes.

While drone-based photogrammetry can provide major insight into cetacean ecology, several drone systems and protocols are used across the scientific community in these efforts, and no consistent method or centralized framework is established for quantifying and incorporating measurement uncertainty associated with these different drones. This lack of standardization restricts comparability across datasets, thus hindering our ability to effectively monitor populations and understand the drivers of variation (e.g., pollution, climate change, injury, noise).

We are excited to announce the Marine Mammal Institute’s (MMI) Center of Drone Excellence (CODEX), which focuses on developing analytical methods for using drones to non-invasively monitor marine mammal populations. CODEX is led by GEMM Lab member’s KC Bierlich, Leigh Torres, and Clara Bird and consists of other team members within and outside OSU. We draw from many years of trials, errors, headaches, and effort working with drones to study cetacean ecology in a variety of habitats and conditions on many different species.

Already CODEX has developed several open-source hardware and software tools. We developed, produced, and published LidarBoX (Bierlich et al., 2023), which is a 3D printed enclosure for a LiDAR altimeter system that can be easily attached and swapped between commercially available drones (i.e., DJI Inspire, DJI Mavic, and DJI Phantom) (Figure 1). Having a LidarBoX installed helps researchers obtain altitude readings with greater accuracy, yielding morphological measurements with less uncertainty. Since we developed LidarBoX, we have received over 35 orders to build this unit for other labs in national and international universities.

Figure 1. A ‘LidarBoX’ attached to a DJI Inspire 2. The LidarBoX is a 3D printed enclosure containing a LiDAR altimeter to help obtain more accurate altitude readings.

Additionally, CODEX recently released MorphoMetriX version 2 (v2), an easy-to-use photogrammetry software that provides users with the flexibility to obtain custom morphological measurements of megafauna in imagery with no knowledge of any scripting language (Torres and Bierlich, 2020). CollatriX is a user-friendly software for collating multiple MorphoMetriX outputs into a single dataframe and linking important metadata to photogrammetric measurements, such as altitude measured with a LidarBoX (Bird and Bierlich, 2020). CollatriX also automatically calculates several body condition metrics based on measurements from MorphoMetriX v2. CollatriX v2 is currently in beta-testing and scheduled to be released late Spring 2024. 

Figure 2. An example of a Pygmy blue whale imported into MorphoMetriX v2, open-source photogrammetry software. 

CODEX also recently developed two automated tools to help speed up the laborious manual processing of drone videos for obtaining morphological measurements (Bierlich & Karki et al., in revision). DeteX is a graphical user interface (GUI) that uses a deep learning model for automated detection of cetaceans in drone-based videos. Researchers can input their drone-based videos and DeteX will output frames containing whales at the surface. Users can then select which frames they want to use for measuring individual whales and then input these selected frames into XtraX, which is a GUI that uses a deep learning model to automatically extract body length and body condition measurements of cetaceans (Figure 4). We found automated measurements from XtraX to be similar (within 5%) of manual measurements. Importantly, using DeteX and XtraX takes about 10% of the time it would take to manually process the same videos, demonstrating how these tools greatly speed up obtaining key morphological data while maintaining accuracy, which is critical for effectively monitoring population health.

Figure 3. An example of an automated body length (top) and body condition (bottom) measurement of a gray whale using XtraX (Bierlich & Karki et al., in revision).

CODEX is also in the process of developing Xcertainty, an R package that uses a Bayesian statistical model to quantify and incorporate uncertainty associated with measurements from different drones (see this blog). Xcertainty is based on the Bayesian statistical model developed by Bierlich et al., (2021b; 2021a), which has been utilized by many studies with several different drones to compare body condition and body morphology across individuals and populations  (Bierlich et al., 2022; Torres et al., 2022; Barlow et al., 2023). Rather than a single point-estimate of a length measurement for an individual, Xcertainty produces a distribution of length measurements for an individual so that the length of a whale can be described by the mean of this distribution, and its uncertainty as the the variance or an interval around the mean (Figure 4). These outputs ensure measurements are robust and comparable across different drones because they provide a measure of the uncertainty around each measurement. For instance, a measurement with more uncertainty will have a wider distribution. The uncertainty associated with each measurement can be incorporated into analyses, which is key when detecting important differences or changes in individuals or populations, such as changes in body condition (blog).

Figure 4. An example of a posterior predictive distribution for total length of an individual blue whale produced by the ‘Xcertainty’ R package. The black bars represent the uncertainty around the mean value (the black dot) – the longer black bars represent the 95% highest posterior density (HPD) interval, and the shorter black bars represent the 65% HPD interval. 

CODEX has integrated all these lessons learned, open-source tools, and analytical approaches into a single framework of suggested best practices to help researchers enhance the quality, speed, and accuracy of obtaining important morphological measurements to manage vulnerable populations. These tools and frameworks are designed to be accommodating and accessible to researchers on various budgets and to facilitate cross-lab collaborations. CODEX plans to host workshops to educate and train researchers using drones on how to apply these tools within this framework within their own research practices. Potential future directions for CODEX include developing a system for using drones to drop suction-cup tags on whales and to collect thermal imagery of whales for health assessments. Stay up to date with all the CODEX ‘X’travaganza here: https://mmi.oregonstate.edu/centers-excellence/codex.  

Huge shout out to Suzie Winquist for designing the artwork for CODEX!

References

Barlow, D.R., Bierlich, K.C., Oestreich, W.K., Chiang, G., Durban, J.W., Goldbogen, J.A., Johnston, D.W., Leslie, M.S., Moore, M.J., Ryan, J.P. and Torres, L.G., 2023. Shaped by Their Environment: Variation in Blue Whale Morphology across Three Productive Coastal Ecosystems. Integrative Organismal Biology, [online] 5(1). https://doi.org/10.1093/iob/obad039.

Bierlich, K., Karki, S., Bird, C.N., Fern, A. and Torres, L.G., n.d. Automated body length and condition measurements of whales from drone videos for rapid assessment of population health. Marine Mammal Science.

Bierlich, K.C., Hewitt, J., Bird, C.N., Schick, R.S., Friedlaender, A., Torres, L.G., Dale, J., Goldbogen, J., Read, A.J., Calambokidis, J. and Johnston, D.W., 2021a. Comparing Uncertainty Associated With 1-, 2-, and 3D Aerial Photogrammetry-Based Body Condition Measurements of Baleen Whales. Frontiers in Marine Science, 8. https://doi.org/10.3389/fmars.2021.749943.

Bierlich, K.C., Hewitt, J., Schick, R.S., Pallin, L., Dale, J., Friedlaender, A.S., Christiansen, F., Sprogis, K.R., Dawn, A.H., Bird, C.N., Larsen, G.D., Nichols, R., Shero, M.R., Goldbogen, J., Read, A.J. and Johnston, D.W., 2022. Seasonal gain in body condition of foraging humpback whales along the Western Antarctic Peninsula. Frontiers in Marine Science, 9(1036860), pp.1–16. https://doi.org/10.3389/fmars.2022.1036860.

Bierlich, K.C., Schick, R.S., Hewitt, J., Dale, J., Goldbogen, J.A., Friedlaender, A.S. and Johnston, D.W., 2021b. Bayesian approach for predicting photogrammetric uncertainty in morphometric measurements derived from drones. Marine Ecology Progress Series, 673, pp.193–210. https://doi.org/10.3354/meps13814.

Bird, C. and Bierlich, K.C., 2020. CollatriX: A GUI to collate MorphoMetriX outputs. Journal of Open Source Software, 5(51), pp.2323–2328. https://doi.org/10.21105/joss.02328.

Torres, L.G., Bird, C.N., Rodríguez-González, F., Christiansen, F., Bejder, L., Lemos, L., Urban R, J., Swartz, S., Willoughby, A., Hewitt, J. and Bierlich, K.C., 2022. Range-Wide Comparison of Gray Whale Body Condition Reveals Contrasting Sub-Population Health Characteristics and Vulnerability to Environmental Change. Frontiers in Marine Science, 9(April), pp.1–13. https://doi.org/10.3389/fmars.2022.867258.

Torres, W. and Bierlich, K.C., 2020. MorphoMetriX: a photogrammetric measurement GUI for morphometric analysis of megafauna. Journal of Open Source Software, 5(45), pp.1825–1826. https://doi.org/10.21105/joss.01825.

A smaller sized gray whale: recent publication finds PCFG whales are smaller than ENP whales

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

A recent blog post by GEMM Lab’s PhD Candidate Clara Bird gave a recap of our 8th consecutive GRANITEfield season this year. In her blog, Clara highlighted that we saw 71 individual gray whales this season, 61 of which we have seen in previous years and identified as belonging to the Pacific Coast Feeding Group (PCFG). With an estimated population size of around 212 individuals, this means that we saw almost 1/3 of the PCFG population this season alone. Since the GEMM Lab first started collecting data on PCFG gray whales in 2016, we have collected drone imagery on over 120 individuals, which is over half the PCFG population. This dataset provides incredible opportunity to get to know these individuals and observe them from year to year as they grow and mature through different life history stages, such as producing a calf. A question our research team has been interested in is what makes a PCFG whale different from an Eastern North Pacific (ENP) gray whale, which has a population size around 16,000 individuals and feed predominantly in the Arctic during the summer months? For this blog, I will highlight findings from our recent publication in Biology Letters (Bierlich et al., 2023) comparing the morphology (body length, skull, and fluke size) between PCFG and ENP populations. 

Body size and shape reflect how an animal functions in their environment and can provide details on an individual’s current health, reproductive status, and energetic requirements. Understanding how animals grow is a key component for monitoring the health of populations and their vulnerability to climate change and other stressors in their environment.  As such, collecting accurate morphological measurements of individuals is essential to model growth and infer their health. Collecting such morphological measurements of whales is challenging, as you cannot ask a whale to hold still while you prepare the tape measure, but as discussed in a previous blog, drones provide a non-invasive method to collect body size measurements of whales. Photogrammetry is a non-invasive technique used to obtain morphological measurements of animals from photographs. The GEMM Lab uses drone-based photogrammetry to obtain morphological measurements of PCFG gray whales, such as their body length, skull length (as snout-to-blowhole), and fluke span (see Figure 1). 

Figure 1. Morphological measurements obtained via photogrammetry of a Pacific Coast Feeding Group (PCFG) gray whale. These measurements were used to compare to individuals from the Eastern North Pacific (ENP) population. 

As mentioned in this previous blog, we use photo-identification to identify unique individual gray whales based on markings on their body. This method is helpful for linking all the data we are collecting (morphology, hormones, behavior, new scarring and skin conditions, etc.) to each individual whale. An individual’s sightings history can also be used to estimate their age, either as a ‘minimum age’ based on the date of first sighting or a ‘known age’ if the individual was seen as a calf. By combining the length measurements from drone-based photogrammetry and age estimates from photo-identification history, we can construct length-at-age growth models to examine how PCFG gray whales grow. While no study has previously examined length-at-age growth models specifically for PCFG gray whales, another study constructed growth curves for ENP gray whales using body length and age estimates obtained from whaling, strandings, and aerial photogrammetry (Agbayani et al., 2020). For our study, we utilized these datasets and compared length-at-age growth, snout-to-blowhole length, and fluke span between PCFG and ENP whales. We used Bayesian statistics to account and incorporate the various levels of uncertainty associated with data collected (i.e., measurements from whaling vs. drone, ‘minimum age’ vs. ‘known age’). 

We found that while both populations grow at similar rates, PCFG gray whales reach smaller adult lengths than ENP. This difference was more extreme for females, where PCFG females were ~1 m (~3 ft) shorter than ENP females and PCFG males were ~0.5 m (1.5 ft) shorter than ENP males (Figure 2, Figure 3). We also found that ENP males and females have slightly larger skulls and flukes than PCFG male and females, respectively. Our results suggest PCFG whales are shaped differently than ENP whales (Figure 3)! These results are also interesting in light of our previous published study that found PCFG whales are skinnier than ENP whales (see this previous blog post). 

Figure 2. Growth curves (von Bertalanffy–Putter) for length-at-age comparing male and female ENP and PCFG gray whales (shading represents 95% highest posterior density intervals). Points represent mean length and median age. Vertical bars represent photogrammetric uncertainty. Dashed horizontal lines represent uncertainty in age estimates.

Figure 3. Schematic highlighting the differences in body size between Pacific Coast Feeding Group (PCFG) and Eastern North Pacific (ENP) gray whales. 

Our results raise some interesting questions regarding why PCFG are smaller: Is this difference in size and shape normal for this population and are they healthy? Or is this difference a sign that they are stressed, unhealthy and/or not getting enough to eat? Larger individuals are typically found at higher latitudes (this pattern is called Bergmann’s Rule), which could explain why ENP whales are larger since they feed in the Arctic. Yet many species, including fish, birds, reptiles, and mammals, have experienced reductions in body size due to changes in habitat and anthropogenic stressors (Gardner et al., 2011). The PCFG range is within closer proximity to major population centers compared to the ENP foraging grounds in the Arctic, which could plausibly cause increased stress levels, leading to decreased growth. 

The smaller morphology of PCFG may also be related to the different foraging tactics they employ on different prey and habitat types than ENP whales. Animal morphology is linked to behavior and habitat (see this blogpost). ENP whales feeding in the Arctic generally forage on benthic amphipods, while PCFG whales switch between benthic, epibenthic and planktonic prey, but mostly target epibenthic mysids. Within the PCFG range, gray whales often forage in rocky kelp beds close to shore in shallow water depths (approx. 10 m) that are on average four times shallower than whales feeding in the Arctic. The prey in the PCFG range is also found to be of equal or higher caloric value than prey in the Arctic range (see this blog), which is interesting since PCFG were found to be skinnier.

It is also unclear when the PCFG formed? ENP and PCFG whales are genetically similar, but photo-identification history reveals that calves born into the PCFG usually return to forage in this PCFG range, suggesting matrilineal site fidelity that contributes to the population structure. PCFG whales were first documented off our Oregon Coast in the 1970s (Figure 4). Though, from examining old whaling records, there may have been PCFG gray whales foraging off the coasts of Northern California to British Columbia since the 1920s.

Figure 4. First reports of summer-resident gray whales along the Oregon coast, likely part of the Pacific Coast Feeding Group. Capital Journal, August 9, 1976, pg. 2.

Altogether, our finding led us to two hypotheses: 1) the PCFG range provides an ecological opportunity for smaller whales to feed on a different prey type in a shallow environment, or 2) the PCFG range is an ecological trap, where individuals gain less energy due to energetically costly feeding behaviors in complex habitat while potentially targeting lower density prey, causing them to be skinnier and have decreased growth. Key questions remain for our research team regarding potential consequences of the smaller sized PCFG whales, such as does the smaller body size equate to reduced resilience to environmental and anthropogenic stressors? Does smaller size effect fecundity and population fitness? Stay tuned as we learn more about this unique and fascinating smaller sized gray whale. 

References

Agbayani, S., Fortune, S. M. E., & Trites, A. W. (2020). Growth and development of North Pacific gray whales (Eschrichtius robustus). Journal of Mammalogy101(3), 742–754. https://doi.org/10.1093/jmammal/gyaa028

Bierlich, K. C., Kane, A., Hildebrand, L., Bird, C. N., Fernandez Ajo, A., Stewart, J. D., Hewitt, J., Hildebrand, I., Sumich, J., & Torres, L. G. (2023). Downsized: gray whales using an alternative foraging ground have smaller morphology. Biology Letters19(8). https://doi.org/10.1098/rsbl.2023.0043

Gardner, J. L., Peters, A., Kearney, M. R., Joseph, L., & Heinsohn, R. (2011). Declining body size: A third universal response to warming? Trends in Ecology and Evolution26(6), 285–291. https://doi.org/10.1016/j.tree.2011.03.005

Title: “Blown away”: measuring the blowholes of whales from drones

By Annie Doron, Undergraduate Intern, Oregon State University, GEMM Laboratory  

Hey up! My name is Annie Doron, and I am an undergraduate Environmental Science student from the University of Sheffield (UK) on my study year abroad. One of my main motivations for undertaking this year abroad was to gain experience working in a marine megafauna lab. Whales in particular have always captivated my interest, and I have been lucky enough to observe  humpback whales in Iceland and The Azores, and even encountered one whilst diving in Australia! For the past 10 months, I have had the unique opportunity to work in the GEMM Lab analyzing Pacific Coast Feeding Group (PCFG) gray whales off the Oregon Coast (Figure 1). I must admit, it has been simply wonderful! 

Figure 1. Aerial image of a PCFG gray whale off the Oregon Coast. 

How did I end up getting involved with the GEMM Lab? I was first accepted into Scarlett Arbuckle’s research-based class in fall term 2022, which is centered around partnering with a mentor for a research project. Having explored the various fields of research at HMSC, I contacted Leigh Torres with interest in getting involved in the GEMM Lab and to establish a research project suitable for a totally inexperienced, international, undergraduate student. Thankfully, Leigh forwarded my email to KC Bierlich who offered to be my mentor for the class, and the rest is history! I first began analyzing drone imagery to measure length and body condition of  PCFG gray whales, which provided an opportunity to get involved with the lab and gain experience using the photogrammetry software MorphoMetriX (Torres & Bierlich, 2020) (see KC’s blog), which is used to make morphometric measurements of whales. Viewing drone imagery of whales sparked my interest in how they use their blowholes (otherwise called ‘nares’) to replenish their oxygen stores; this led to us establishing a research project for the class where we tested if we could use MorphoMetriX to measure blowholes from drone imagery.

Extending this project into winter and spring terms (via research credits) has enabled me to continue working with Leigh and KC, as well as to collaborate with Clara Bird and Jim Sumich. Thanks to KC, who has patiently guided me through the ins and outs of working on a research project, I now feel more confident handling and manipulating large datasets, analyzing drone footage (i.e., differentiating between behavioral states, recording breathing sequences, detecting when a whale is exhaling vs inhaling, etc.), and speaking in public (although I still get pretty bad stage fright, but I think that is a typical conundrum undergrads face). Whatsmore, applying  R – a programming language used for statistical analysis and data visualization, which I have been trying to wrap my head around for years – to my own dataset has helped me greatly enhance my skills using it. 

So, what exciting things have we been working on this year? Given that we often cannot simply study a whale from inside a laboratory – due to size-related logistical implications – we must use proxies (i.e., a variable that is representative of an immeasurable variable). Since cetaceans must return to the surface to offload carbon dioxide and replenish their oxygen stores, measuring their breath frequency and magnitude is one way to study a whale’s oxygen consumption, in turn offering insight into its energy expenditure (Williams, 1999). Blowholes are one proxy we can use to study breath magnitude. Blowholes can be utilized in this way by measuring inhalation duration (the amount of time a whale is inhaling, which is based on a calculation developed by Jim Sumich) and blowhole area (the total area of a blowhole) to gauge variations in tidal volume (the amount of air flowing in and out of the lungs).

Measuring inhalation duration and blowhole area is important because a larger blowhole area (i.e., one that is more dilated) and a longer inhalation duration is indicative of higher oxygen intake, which can infer stress. For example, in this population, higher stress levels are associated with increased vessel traffic (Lemos et al., 2022), and skinnier whales have higher stress levels compared to chubby, healthy whales (Lemos, Olsen, et al., 2022). Hence, measuring the variation around blowholes could be utilized to predict challenges whales face from climate change and anthropogenic disturbance, including fishing (Scordino et al., 2017) and whale watching industry threats (Sullivan & Torres, 2018) (see Clara’s blog), as well as to inform effective management strategies. Furthermore, measuring the variables inhalation duration and blowhole area could help to identify whether whales are taking larger breaths associated with certain ‘gross behavior states’, otherwise known as ‘primary states’, which include: travel, forage, rest, social (Torres et al., 2018). This could enable us to assess the energetic costs of different foraging tactics (i.e., head standing, side-swimming, and bubble blasting (Torres et al., 2018), as well as consequences of disturbance events, on an individual and population health perspective. 

Inhalation duration has been explored in the past by using captive animals. For example, there have been studies on heart rate and breathing of bottlenose dolphins in human care facilities (Blawas et al., 2021; Fahlman et al., 2015). Recently, Nazario et al. (2022) was able to measure inhalation duration and blowhole area using suction-cup video tags. Her study led us to consider if it was possible to measure the parameters and variation around respiration by measuring blowhole area and inhalation duration of PCFGs from drone imagery. We employed MorphoMetriX to study the length, width, and area of a blowhole (Figure 2). Preliminary analyses verified that the areas of the left and right blowholes are very similar (Figure 3); this finding saved us a lot of time because from thereon we only measured either the left or right side. Interestingly, we see some variation in blowhole area within and across individuals (Figure 4). This variation changes within individuals based on primary state. For example, the whales “Glacier”, “Nimbus”, and “Rat” show very little variation whilst traveling but a large amount whilst foraging. Comparatively, “Dice” shows little variation whilst foraging and large variation whilst traveling. Whilst considering cross-individual comparisons, we can see that “Sole”, “Rat”, “Nimbus”, “Heart”, “Glacier”, “Dice”, and “Coal” each exhibit relatively large amounts of variation, yet “Mahalo”, “Luna”, “Harry”, “Hummingbird” and “Batman” exhibit very little. One potential reason for some individuals displaying higher levels of variation than others could be higher levels of exposure to disturbance events that we were unable to measure or evaluate in this study.

Figure 2. How we measured the length, width, and area of a blowhole using MorphoMetriX.

Figure 3. Data driven evidence that the left and the right blowhole areas are very similar. 

Figure 4. Variation in blowhole area amongst individual PCFG whales. The hollow circles represent the means, and the color represents the primary state the whale is exhibiting, foraging (purple) vs. traveling (blue), which will be further explored in Clara’s PhD.

Now, we are venturing into June and are at a stage where we (KC, Clara, Jim, Leigh, and I) are preparing to publish a manuscript! What a way to finish such a fantastic year! The transition from a 3-month-long pilot study to a much larger data analysis and eventual preparation for a manuscript has been a monumental learning experience. If anybody had told me a year ago that I would be involved in publishing a body of work – especially one that is so meaningful to me – I would simply not have believed them! We hope this established methodology for measuring blowholes will help other researchers carry out blowhole measurements using drone imagery across different populations and species. Further research is required to explore the differences in inhalation duration and blowhole area between different primary states, specifically across different foraging tactics.

It has been a great privilege working with the GEMM Lab these past months, and I was grateful to be included in their monthly lab meetings, during which members gave updates and we discussed recently published papers. Seeing such an enthusiastic, kind, and empathic group of people working together taught me what working in a supportive lab could look and feel like. In spite of relocating from Corvallis to Bend after my first term, I was happy to be able to continue working remotely for the lab for the remainder of my time (even though I was ~200 miles inland). I thoroughly enjoyed living in Corvallis, highlights of which were scuba diving adventures to the Puget Sound and coastal road trips with friends. The appeal to move arose from Bend’s reputation as an adventure hub – with unlimited opportunities for backcountry ski access – as well as its selection of wildlife ecology courses (with a focus on species specific to central Oregon). I moved into ‘Bunk & Brew’ (Bend’s only hostel, which is more like a big house of friends with occasional hostel guests) on January 1st after returning from spending Christmas with friends in my old home in Banff, Canada. I have since been enjoying this wonderful multifaceted lifestyle; working remotely in the GEMM Lab, attending in-person classes, working part-time at the hostel, as well as skiing volcanoes (Mount Hood, Middle and South Sister (Figure 5) or climbing at Smith Rock during my days off. Inevitably, I do miss the beautiful Oregon coast, and I will always be grateful for this ideal opportunity and hope this year marks the start of my marine megafauna career!

Figure 5. What I get up to when I’m not studying blowholes! (This was taken at 5am on the long approach to Middle and North Sister. North Sister is the peak featured in the backdrop).

References

Blawas, A. M., Nowacek, D. P., Allen, A. S., Rocho-Levine, J., & Fahlman, A. (2021). Respiratory sinus arrhythmia and submersion bradycardia in bottlenose dolphins (Tursiops truncatus). Journal of Experimental Biology, 224(1), jeb234096. https://doi.org/10.1242/jeb.234096

Fahlman, A., Loring, S. H., Levine, G., Rocho-Levine, J., Austin, T., & Brodsky, M. (2015). Lung mechanics and pulmonary function testing in cetaceans. Journal of Experimental Biology, 218(13), 2030–2038. https://doi.org/10.1242/jeb.119149

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), 18580. https://doi.org/10.1038/s41598-022-14510-5

Lemos, L. S., Olsen, A., Smith, A., Burnett, J. D., Chandler, T. E., Larson, S., Hunt, K. E., & Torres, L. G. (2022). Stressed and slim or relaxed and chubby? A simultaneous assessment of gray whale body condition and hormone variability. Marine Mammal Science, 38(2), 801–811. https://doi.org/10.1111/mms.12877

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

Scordino, J., Carretta, J., Cottrell, P., Greenman, J., Savage, K., & Scordino, J. (2017). Ship Strikes and Entanglements of Gray Whales in the North Pacific Ocean. Cambridge: International Whaling Commission, 1924–2015.

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

Sumich, J. L. (1994). Oxygen extraction in free-swimming gray whale caves. Marine Mammal Science, 10(2), 226–230. https://doi.org/10.1111/j.1748-7692.1994.tb00266.x

Torres, W., & Bierlich, K. (2020). MorphoMetriX: A photogrammetric measurement GUI for morphometric analysis of megafauna. Journal of Open Source Software, 5(45), 1825. https://doi.org/10.21105/joss.01825

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. https://doi.org/10.3389/fmars.2018.00319
Williams, T. M. (1999). The evolution of cost efficient swimming in marine mammals: Limits to energetic optimization. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 354(1380), 193–201. https://doi.org/10.1098/rstb.1999.0371

So big, but so small: why the smallest of the largest whales are not smaller

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

Baleen whales are known for their gigantism and encompass a wide range in body sizes extending from blue whales that are the largest animals to live on earth (max length ~30 m) to minke whales (max length ~10 m) that are the smallest of baleen whales (Fig. 1). While all baleen whales are filter feeders, a group called the rorquals use a feeding strategy known as lunge feeding (or intermittent engulfment filtration), which involves engulfing large volumes of prey-laden water at high speeds and then filtering the water out of their mouth using their baleen as a “sieve”. There is positive allometry associated with this feeding technique and body size, meaning that as whales are larger, this feeding strategy becomes more efficient due to increased engulfment of water volume per each lunge feeding event. In other words, a bigger body size equates to a much larger mouthful of food. For example, a minke whale (body length ~7-10 m) will engulf water volume equivalent to ~42% of its body mass, while a blue whale (~21-24 m) engulfs ~135%. Thus, filter feeding enables gigantism through efficient exploitation of large, dense patches of prey. An interesting question then arises: what is the minimum body size at which filter feeding is still efficient? Or in other words, why are the smallest of the baleen whales, minke whales, not smaller? For this blog, I will highlight a study published today in Nature Ecology and Evolution titled “Minke whale feeding rate limitations suggest constraints on the minimum body size for engulfment filtration feeding” led by friend and collaborator of the GEMM Lab Dr. Dave Cade and included myself and other collaborators as co-authors from Stanford University, UC Santa Cruz, Cascadia Research Collective, Duke University, and University of Queensland.

Figure 1. Aerial imagery collected using drones of several baleen whales of various sizes. Each species shown is considered a rorqual whale, except for gray whales. Figure from Segre et al. (2022)

The largest animals of today are marine filter feeders, such as whale sharks, manta rays, and baleen whales, which all share parallel evolutionary histories in which their large body sizes and filter-feeding morphologies are derived from smaller-bodied ancestors that targeted single prey items. Changes in ocean productivity increased the concentrations of smaller prey in the oceans around 5 million years ago, enabling filter feeding as an efficient feeding strategy through capture of abundant aggregations of prey by filtering large volumes of water. It is interesting to note, that within these filter feeding lineages of animals, there are groups of animals that are single-prey foragers with smaller body sizes. For example, the whale shark is the only filter feeder amongst the carpet sharks and the manta ray is much larger than other rays that feed on single prey items. Amongst cetaceans, the smallest single-prey foragers, dolphins (~2-3 m) and porpoises (~1.4-1.9 m), are much smaller than the smallest of the filter feeding cetaceans, minke whales (~7-10 m). These common differences in body sizes and feeding strategies within lineages suggest that there may be minimum body size requirements for this filter feeding strategy to be efficient.

To investigate the limits on minimum body size for filter feeding, our study explored the foraging behavior of Antarctic minke whales, the smallest of the rorqual baleen whales, along the Western Antarctic Peninsula. Our team tagged a total of 23 individuals using non-invasive suction cup tags, like the ones we use for our tagging component in the GEMM Lab’s GRANITE project (see this blog for more details). One of my roles on the project was to obtain aerial imagery of the minke whales using drones to obtain body length measurements (sound familiar?) (Figs. 2-4). Flying drones in Antarctica over minke whales was an amazing experience. The minke whales were often found deep within the bays amongst ice floes and brash ice where they can be very tricky to spot, as they’ll often surface and then quickly disappear, hence their nickname “sneaky minkes”. They also appear “playful” and “athletic” as they are incredibly quick and maneuverable, doing barrel rolls and quick bank turns while they swim. Check out my past blog to read more on accounts of flying over these amazing whales.

Figure 2. Drone image of our team about to place a noninvasive suction cup biologging tag on an Antarctic minke whale. Photo credit: Duke University Marine Robotics and Remote Sensing Lab.
Figure 3. A drone image of a newly tagged and curious Antarctic minke whale approaching our research team. Photo credit: Duke University Marine Robotics and Remote Sensing Lab.
Figure 4. A drone image of a group of Antarctic minke whales swimming through the icy waters along the Antarctic Peninsula. Photo credit: Duke University Marine Robotics and Remote Sensing Lab.

In total, our team collected 437 hours of tag data consisting of day- and night-time foraging behaviors. While the proportion of time spent foraging and the number of lunges per dive (~3-4) was similar between day- and night-time foraging, daytime foraging was much deeper (~72 m) compared to nighttime foraging (~28 m) due to vertical migration of Antarctic krill, their main food source. Overall, nighttime foraging was much more intense than daytime foraging, with an average of 165 lunges per hour during the night compared to 53 lunges per hour during the day. These shallower nighttime dives enabled quicker surface sequences for replenishing oxygen reserves to then return to foraging, whereas the deeper dives during the day required longer surface recovery times before beginning another foraging dive. Thus, nighttime dives are a more efficient and critical component of minke whale foraging.

When it comes to body size, there was no relationship between dive depth and dive duration with body length, except for daytime deep dives, where longer minke whales dove for longer periods than smaller whales. These longer dive times also require longer surface times to replenish oxygen reserves. Longer minke whales can gulp larger amounts of food and thus need longer filtration times to process water from each engulfment. For example, a 9 m minke whale will take 50% longer to filter water through its baleen compared to a 5 m minke whale. In turn, smaller minke whales would need to feed more frequently than larger minke whales in order to maintain efficient foraging. This decreasing efficiency with smaller body size shines light on a broader trend for filter feeders that we refer to in our study as the minimum-size constraint (MSC) hypothesis: “while the maximum size of a filter-feeding body plan will be restricted by physical properties, the minimum size is restricted by the energetic efficiency of filter feeding and the time required to extract sufficient particles from the water” (Cade et al. 2023). When we examined the scaling of maximum feeding rates of minke whales, we found evidence of a minimum size constraint on efficiency at lengths around 5 m. Interestingly, the weaning length of minke whales is reported to be 4.5 – 5.5 m. Before weaning, newborn/yearling minke whales that are smaller than 4.5 ­– 5.5 m have a different foraging strategy where they are dependent on maternal milk. Thus, it is likely that the body size at weaning is influenced by the minimum size at which this specialized foraging technique of lunge feeding becomes efficient.

This study helps inform the evolutionary pathway for filter feeding whales and suggests that efficient filter feeding and gigantism likely co-evolved within the last 5 million years when ocean conditions changed to support larger prey patches suitable for lunge feeding. It is interesting to think about the MSC hypothesis for other baleen whale species that employ alternative filter feeding techniques, such as gray whales that generally use a form of filter feeding called suction feeding. Gray whales are estimated to have a birth length of ~4.6 m (Agbayani et al., 2020), and the body length of newly weaned calves that we have observed along the Oregon Coast from drone imagery seem to be ~8 – 9 m. Perhaps this is the minimum size of when suction feeding becomes efficient for a gray whale? This is something the GEMM Lab hopes to further explore as we continue to collect foraging data from suction cup tags and behavior and body size measurements from drone imagery.

References

Agbayani, S., Fortune, S. M., & Trites, A. W. (2020). Growth and development of North Pacific gray whales (Eschrichtius robustus). Journal of Mammalogy101(3), 742-754.

Cade, D.E., Kahane-Rapport, S.R., Gough, W.T., Bierlich, K.C., Linksy, J.M.J., Johnston, D.W., Goldbogen, J.A., Friedlaender, A.S. (2023). Ultra-high feeding rates of Antarctic minke whales imply a lower limit for body size in engulfment filtration feeders. Nature Ecology and Evolution. https://www.nature.com/articles/s41559-023-01993-2  

Paolo S. Segre, William T. Gough, Edward A. Roualdes, David E. Cade, Max F. Czapanskiy, James Fahlbusch, Shirel R. Kahane-Rapport, William K. Oestreich, Lars Bejder, K. C. Bierlich, Julia A. Burrows, John Calambokidis, Ellen M. Chenoweth, Jacopo di Clemente, John W. Durban, Holly Fearnbach, Frank E. Fish, Ari S. Friedlaender, Peter Hegelund, David W. Johnston, Douglas P. Nowacek, Machiel G. Oudejans, Gwenith S. Penry, Jean Potvin, Malene Simon, Andrew Stanworth, Janice M. Straley, Andrew Szabo, Simone K. A. Videsen, Fleur Visser, Caroline R. Weir, David N. Wiley, Jeremy A. Goldbogen; Scaling of maneuvering performance in baleen whales: larger whales outperform expectations. J Exp Biol 1 March 2022; 225 (5): jeb243224. doi: https://doi.org/10.1242/jeb.243224

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

New publication by GEMM Lab reveals sub-population health differences in gray whales 

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

In a previous blog, I discussed the importance of incorporating measurement uncertainty in drone-based photogrammetry, as drones with different sensors, focal length lenses, and altimeters will have varying levels of measurement accuracy. In my last blog, I discussed how to incorporate photogrammetric uncertainty when combining multiple measurements to estimate body condition of baleen whales. In this blog, I will highlight our recent publication in Frontiers in Marine Science (https://doi.org/10.3389/fmars.2022.867258) led by GEMM Lab’s Dr. Leigh TorresClara Bird, and myself that used these methods in a collaborative study using imagery from four different drones to compare gray whale body condition on their breeding and feeding grounds (Torres et al., 2022).

Most Eastern North Pacific (ENP) gray whales migrate to their summer foraging grounds in Alaska and the Arctic, where they target benthic amphipods as prey. A subgroup of gray whales (~230 individuals) called the Pacific Coast Feeding Group (PCFG), instead truncates their migration and forages along the coastal habitats between Northern California and British Columbia, Canada (Fig. 1). Evidence from a recent study lead by GEMM Lab’s Lisa Hildebrand (see this blog) found that the caloric content of prey in the PCFG range is of equal or higher value than the main amphipod prey in the Arctic/sub-Arctic regions (Hildebrand et al., 2021). This implies that greater prey density and/or lower energetic costs of foraging in the Arctic/sub-Arctic may explain the greater number of whales foraging in that region compared to the PCFG range. Both groups of gray whales spend the winter months on their breeding and calving grounds in Baja California, Mexico. 

Figure 1. The GEMM Lab field team following a Pacific Coast Feeding Group (PCFG) gray whale swimming in a kelp bed along the Oregon Coast during the summer foraging season. 

In January 2019 an Unusual Mortality Event (UME) was declared for gray whales due to the elevated numbers of stranded gray whales between Mexico and the Arctic regions of Alaska. Most of the stranded whales were emaciated, indicating that reduced nutrition and starvation may have been the causal factor of death. It is estimated that the population dropped from ~27,000 individuals in 2016 to ~21,000 in 2020 (Stewart & Weller, 2021).

During this UME period, between 2017-2019, the GEMM Lab was using drones to monitor the body condition of PCFG gray whales on their Oregon coastal feeding grounds (Fig. 1), while Christiansen and colleagues (2020) was using drones to monitor gray whales on their breeding grounds in San Ignacio Lagoon (SIL) in Baja California, Mexico. We teamed up with Christiansen and colleagues to compare the body condition of gray whales in these two different areas leading up to the UME. Comparing the body condition between these two populations could help inform which population was most effected by the UME.

The combined datasets consisted of four different drones used, thus different levels of photogrammetric uncertainty to consider. The GEMM Lab collected data using a DJI Phantom 3 Pro, DJI Phantom 4, and DJI Phantom 4 Pro, while Christiansen et al., (2020) used a DJI Inspire 1 Pro. By using the methodological approach described in my previous blog (here, also see Bierlich et al., 2021a for more details), we quantified photogrammetric uncertainty specific to each drone, allowing cross-comparison between these datasets. We also used Body Area Index (BAI), which is a standardized relative measure of body condition developed by the GEMM Lab (Burnett et al., 2018) that has low uncertainty with high precision, making it easier to detect smaller changes between individuals (see blog here, Bierlich et al., 2021b). 

While both PCFG and ENP gray whales visit San Ignacio Lagoon in the winter, we assume that the photogrammetry data collected in the lagoon is mostly of ENP whales based on their considerably higher population abundance. We also assume that gray whales incur low energetic cost during migration, as gray whale oxygen consumption rates and derived metabolic rates are much lower during migration than on foraging grounds (Sumich, 1983). 

Interestingly, we found that gray whale body condition on their wintering grounds in San Ignacio Lagoon deteriorated across the study years leading up to the UME (2017-2019), while the body condition of PCFG whales on their foraging grounds in Oregon concurrently increased. These contrasting trajectories in body condition between ENP and PCFG whales implies that dynamic oceanographic processes may be contributing to temporal variability of prey available in the Arctic/sub-Arctic and PCFG range. In other words, environmental conditions that control prey availability for gray whales are different in the two areas. For the ENP population, this declining nutritive gain may be associated with environmental changes in the Arctic/sub-Arctic region that impacted the predictability and availability of prey. For the PCFG population, the increase in body condition across years may reflect recovery of the NE Pacific Ocean from the marine heatwave event in 2014-2016 (referred to as “The Blob”) that resulted with a period of low prey availability. These findings also indicate that the ENP population was primarily impacted in the die-off from the UME. 

Surprisingly, the body condition of PCFG gray whales in Oregon was regularly and significantly lower than whales in San Ignacio Lagoon (Fig. 2). To further investigate this potential intrinsic difference in body condition between PCFG and ENP whales, we compared opportunistic photographs of gray whales feeding in the Northeastern Chukchi Sea (NCS) in the Arctic collected from airplane surveys. We found that the body condition of PCFG gray whales was significantly lower than whales in the NCS, further supporting our finding that PCFG whales overall have lower body condition than ENP whales that feed in the Arctic (Fig. 3). 

Figure 2. Boxplots showing the distribution of Body Area Index (BAI) values for gray whales imaged by drones in San Ignacio Lagoon (SIL), Mexico and Oregon, USA. The data is grouped by phenology group: End of summer feeding season (departure Oregon vs. arrival SIL) and End of wintering season (arrival Oregon vs. departure SIL). The group median (horizontal line), interquartile range (IQR, box), maximum and minimum 1.5*IQR (vertical lines), and outliers (dots) are depicted in the boxplots. The overlaid points represent the mean of the posterior predictive distribution for BAI of an individual and the bars represents the uncertainty (upper and lower bounds of the 95% HPD interval). Note how PCFG whales at then end of the feeding season (dark green) typically have lower body condition (as BAI) compared to ENP whales at the end of the feeding season when they arrive to SIL after migration (light brown).
Figure 3. Boxplots showing the distribution of Body Area Index (BAI) values of gray whales from opportunistic images collected from a plane in Northeaster Chukchi Sea (NCS) and from drones collected by the GEMM Lab in Oregon. The boxplots display the group median (horizontal line), interquartile range (IQR box), maximum and minimum 1.5*IQR (vertical lines), and outlies (dots). The overlaid points are the BAI values from each image. Note the significantly lower BAI of PCFG whales on Oregon feeding grounds compared to whales feeding in the Arctic region of the NCS.

This difference in body condition between PCFG and ENP gray whales raises some really interesting and prudent questions. Does the lower body condition of PCFG whales make them less resilient to changes in prey availability compared to ENP whales, and thus more vulnerable to climate change? If so, could this influence the reproductive capacity of PCFG whales? Or, are whales that recruit into the PCFG adapted to a smaller morphology, perhaps due to their specialized foraging tactics, which may be genetically inherited and enables them to survive with reduced energy stores?

These questions are on our minds here at the GEMM Lab as we prepare for our seventh consecutive field season using drones to collect data on PCFG gray whale body condition. As discussed in a previous blog by Dr. Alejandro Fernandez Ajo, we are combining our sightings history of individual whales, fecal hormone analyses, and photogrammetry-based body condition to better understand gray whales’ reproductive biology and help determine what the consequences are for these PCFG whales with lower body condition.

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References

Bierlich, K. C., Hewitt, J., Bird, C. N., Schick, R. S., Friedlaender, A., Torres, L. G., … & Johnston, D. W. (2021). Comparing Uncertainty Associated With 1-, 2-, and 3D Aerial Photogrammetry-Based Body Condition Measurements of Baleen Whales. Frontiers in Marine Science, 1729.

Bierlich, K. C., Schick, R. S., Hewitt, J., Dale, J., Goldbogen, J. A., Friedlaender, A.S., et al. (2021b). Bayesian Approach for Predicting Photogrammetric Uncertainty in Morphometric Measurements Derived From Drones. Mar. Ecol. Prog. Ser. 673, 193–210. doi: 10.3354/meps13814

Burnett, J. D., Lemos, L., Barlow, D., Wing, M. G., Chandler, T., & Torres, L. G. (2018). Estimating morphometric attributes of baleen whales with photogrammetry from small UASs: A case study with blue and gray whales. Marine Mammal Science35(1), 108–139.

Christiansen, F., Rodrı́guez-González, F., Martı́nez-Aguilar, S., Urbán, J., Swartz, S., Warick, H., et al. (2021). Poor Body Condition Associated With an Unusual Mortality Event in Gray Whales. Mar. Ecol. Prog. Ser. 658, 237–252. doi:10.3354/meps13585

Hildebrand, L., Bernard, K. S., and Torres, L. G. (2021). Do Gray Whales Count Calories? Comparing Energetic Values of Gray Whale Prey Across Two Different Feeding Grounds in the Eastern North Pacific. Front. Mar. Sci. 8. doi: 10.3389/fmars.2021.683634

Stewart, J. D., and Weller, D. (2021). Abundance of Eastern North Pacific Gray Whales 2019/2020 (San Diego, CA: NOAA/NMFS)

Sumich, J. L. (1983). Swimming Velocities, Breathing Patterns, and Estimated Costs of Locomotion in Migrating Gray Whales, Eschrichtius Robustus. Can. J. Zoology. 61, 647–652. doi: 10.1139/z83-086

Torres, L.G., Bird, C., Rodrigues-Gonzáles, F., Christiansen F., Bejder, L., Lemos, L., Urbán Ramírez, J., Swartz, S., Willoughby, A., Hewitt., J., Bierlich, K.C. (2022). Range-wide comparison of gray whale body condition reveals contrasting sub-population health characteristics and vulnerability to environmental change. Frontiers in Marine Science. 9:867258. https://doi.org/10.3389/fmars.2022.867258

The many dimensions of a fat whale: Using drones to measure the body condition of baleen whales 

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

In my last blog, I discussed how to obtain morphological measurements from drone-based imagery of whales and the importance of calculating and considering uncertainty, as different drone platforms have varying levels of measurement uncertainty. But how does uncertainty scale and propagate when multiple measurements are combined, such as when measuring body condition of the whole animal? In this blog, I will discuss the different methods used for measuring body condition of baleen whales from drone-based imagery and how uncertainty differs between these metrics.

Body condition is defined as the energy stored in the body as a result of feeding and is assumed to indicate an animal’s overall health, as it reflects the balance between energy intake and investment toward growth, maintenance and reproduction (Peig and Green, 2009). Thus, body condition reflects the foraging success of an individual, as well as the potential for reproductive output and the quality of habitat. For example, female North American brown bears (Ursus arctos) in high quality habitats were in better body condition, produced larger litter sizes, and lived in greater population densities compared to females in lower quality habitats (Hilderbrand et al., 1999). As Dawn Barlow and Will Kennerley discussed in their recent blog, baleen whales are top predators and serve as ecosystem sentinels that shed light not only on the health of their population, but on the health of their ecosystem. As ocean climate conditions continue to change, monitoring the body condition of baleen whales is important to provide insight on how their population and ecosystem is responding. 

As discussed in a previous blog, drones serve as a valuable tool for obtaining morphological measurements of baleen whales to estimate their body condition. Images are imported into photogrammetry software, such as MorphoMetriX (Torres and Bierlich, 2020), to measure the total length of an individual and that is then divided into perpendicular width segments (i.e., in 5 or 10% increments) down the body (Fig. 1). These total length and width measurements are then used to estimate body condition in either 1-, 2-, or 3-dimensions: a single width (1D), a projected dorsal surface area (2D), or a body volume measure (3D). These 1D, 2D, and 3D measurements of body condition can then be standardized by total length to produce a relative measure of an individual’s body condition to compare among individuals and populations. 

Figure 1. An example of a Pacific Coast Feeding Group (PCFG) gray whale measured in MorphoMetriX (Torres & Bierlich, 2020).

While several different studies have used each of these dimensions to assess whale body condition, it is unclear how these measurements compare amongst each other. Importantly, it is also unclear how measurement uncertainty scales across these multiple dimensions and influences inference, which can lead to misinterpretation of data. For example, the surface area and volume of two geometrically similar bodies of different sizes are not related to their linear dimensions in the same ratio, but rather to the second and third power, respectively (i.e., x2 vs. x3).  Similarly, uncertainty should not be expected to scale linearly across 1D, 2D, and 3D body condition measurements. 

The second chapter of my dissertation, which was recently published in Frontiers in Marine Science and includes Clara Bird and Leigh Torres as co-authors, compared the uncertainty associated with 1D, 2D, and 3D drone-based body condition measurements in three baleen whale species with different ranges in body sizes: blue, humpback, and Antarctic minke whales (Figure 2) (Bierlich et al., 2021). We used the same Bayesian model discussed in my last blog, to incorporate uncertainty associated with each 1D, 2D, and 3D estimate of body condition. 

Figure 2. An example of total length and perpendicular width (in 5% increments of total length) measurements of an individual blue, humpback and Antarctic minke whale. Each image measured using MorphoMetriX (Torres and Bierlich, 2020). 

We found that uncertainty does not scale linearly across multi-dimensional measurements, with 2D and 3D uncertainty increasing by a factor of 1.45 and 1.76 compared to 1D, respectively. This result means that there is an added cost of increased uncertainty when utilizing a multidimensional body condition measurement. Our finding is important to help researchers decide which body condition measurement best suits their scientific question,  particularly when using a drone platform that is susceptible to greater error – as discussed in my previous blog. However, a 1D measurement only relies on a single width measurement, which may be excluding other regions of an individual’s body condition that is important for energy storage. In these situations, a 2D or 3D measure may be more appropriate.

We found that when comparing relative measures of body condition (standardized by total length of the individual), each standardized metric was highly correlated with one another. This finding suggests that 1D, 2D, and 3D metrics will draw similar relative predictions of body condition for individuals, allowing researchers to be confident they will draw similar conclusions relating to the body condition of individuals, regardless of which standardized metric they use. However, when comparing the precision of each of these metrics, the body area index (BAI) – a 2D standardized metric – displayed the highest level of precision. This result highlights how BAI can advantageously detect small changes in body condition, which is useful for comparing individuals or even tracking the same individual over time.

BAI was developed by the GEMM Lab (Burnett et al., 2018) and was designed to be similar to body mass index (BMI) in humans [BMI = mass (kg)/(height (m))2], where BAI uses the calculated surface area as a surrogate for body mass. In humans, a healthy BMI range is generally considered 18.5–24.9, below 18.5 is considered underweight, above 24.9 is considered overweight, and above 30 is considered obese (Flegal et al., 2012). Identifying a healthy range in BAI for baleen whales is challenged by a limited knowledge of what a “healthy” body condition range is for a whale. We found strong evidence that a healthy range of BAI is species-specific, as each species displayed a distinctive range in BAI: blue whales: 11–16; AMW: 17–24; humpback whales: 23–32; humpback whale calves: 23–28 (Fig. 3). These differences in BAI ranges likely reflect differences in the body shape of each species (Fig. 4). For example, humpbacks have the widest range of BAI compared to these other two species, which was also reflected in their larger variation in perpendicular widths (Figs. 2-4). Thus, it seems that BAI offers conditionally “scalefree” comparisons between species, yet it is unreasonable to set a single, all-whale BAI threshold to determine “healthy” versus “unhealthy” body condition.  Collecting a large sample of body condition measurements across many individuals and demographic units over space and time with information on vital rates (e.g., reproductive capacity) will help elucidate a healthy BAI range for each species.

Figure 3. Body area index (BAI) for each species. AMW = Antarctic minke whale.  Figure from Bierlich et al. (2021).
Figure 4. A) Absolute widths (m) and B) relative widths, standardized by total length (TL) to help elucidate the different body shapes of Antarctic minke whales (AMW; n = 40), blue whales (n = 32), humpback whales (n = 40), and humpback whale calves (n = 15). Note how the peak in body width occurs at a different percent body width between species, demonstrating the natural variation in body shape between baleen whales. Figure from Bierlich et al. (2021).

Over the past six years, the GEMM Lab has been collecting drone images of Pacific Coast Feeding Group (PCFG) gray whales off the coast of Oregon to measure their BAI (see GRANITE Project blog). Many of the individuals we encounter are seen across years and throughout the foraging season, providing an opportunity to evaluate how an individual’s BAI is influenced by environmental variation, stress levels, maturity, and reproduction. These data will in turn help determine what the healthy range in BAI for gray whales is. For example, linking BAI to pregnancy – whether a whale is currently pregnant or becomes pregnant the following season – will help determine what BAI is needed to support calf production. We are currently analyzing hundreds of body condition measurements from 2016 – 2021, so stay tuned for upcoming results!

References

Bierlich, K. C., Hewitt, J., Bird, C. N., Schick, R. S., Friedlaender, A., Torres, L. G., … & Johnston, D. W. (2021). Comparing Uncertainty Associated With 1-, 2-, and 3D Aerial Photogrammetry-Based Body Condition Measurements of Baleen Whales. Frontiers in Marine Science, 1729.

Burnett, J. D., Lemos, L., Barlow, D., Wing, M. G., Chandler, T., & Torres, L. G. (2018). Estimating morphometric attributes of baleen whales with photogrammetry from small UASs: A case study with blue and gray whales. Marine Mammal Science35(1), 108–139.

Flegal, K. M., Carroll, M. D., Kit, B. K., & Ogden, C. L. (2012). Prevalence of Obesity and Trends in the Distribution of Body Mass Index Among US Adults, 1999-2010. JAMA307(5), 491. https://doi.org/10.1001/jama.2012.39

Hilderbrand, G. V, Schwartz, C. C., Robbins, C. T., Jacoby, M. E., Hanley, T. A., Arthur, S. M., & Servheen, C. (1999). The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Canadian Journal of Zoology77(1), 132–138.

Peig, J., & Green, A. J. (2009). New perspectives for estimating body condition from mass/length data: the scaled mass index as an alternative method. Oikos118(12), 1883–1891.

Torres, W., & Bierlich, K. C. (2020). MorphoMetriX: a photogrammetric measurement GUI for morphometric analysis of megafauna. Journal of Open Source Software5(45), 1825–1826.

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. Neuron84(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 Communications12(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 Neurobiology68, 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 Zoology68(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 Biology30(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 Series629, 219–234. https://doi.org/10.3354/meps13125

Pereira, T. D., Shaevitz, J. W., & Murthy, M. (2020). Quantifying behavior to understand the brain. Nature Neuroscience23(12), 1537–1549. https://doi.org/10.1038/s41593-020-00734-z

The learning curve never stops as the GRANITE project begins its seventh field season

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!

Connecting Research Questions

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

The field season can be quite a hectic time of year. Between long days out on the water, trouble-shooting technology issues, organizing/processing the data as it comes in, and keeping up with our other projects/responsibilities, it can be quite overwhelming and exhausting.

But despite all of that, it’s an incredible and exciting time of year. Outside of the field season, we spend most of our time staring at our computers analyzing the data that we spend a relatively short amount of time collecting. When going through that process it can be easy to lose sight of why we do what we do, and to feel disconnected from the species we are studying. Oftentimes the analysis problems we encounter involve more hours of digging through coding discussion boards than learning about the animals themselves. So, as busy as it is, I find that the field season can be pretty inspiring. I have recently been looking through our most recent drone footage of gray whales and feeling renewed excitement for my thesis.

At the moment, my thesis has four central questions: (1) Are there associations between habitat type and gray whale foraging tactic? (2) Is there evidence of individualization? (3) What is the relationship between behavior and body condition? (4) Do we see evidence of learning in the behavior of mom and calf pairs? As I’ve been organizing my thoughts, what’s become quite clear is how interconnected these questions are. So, I thought I’d take this blog to describe the potential relationships.

Let’s start with the first question: are there associations between habitat types and gray whale foraging tactics? This question is central because it relates foraging behavior to habitat, which is ultimately associated with prey. This relationship is the foundation of all other questions involving foraging tactics because food is necessary for the whales to have the energy and nutrients they need to survive. It’s reasonable to think that the whales are flexible and use different foraging tactics to eat different prey that live in different habitats. But, if different prey types have different nutritional value (this is something that Lisa is studying right now; check out the COZI project to learn more), then not all whales may be getting the same nutrients.

The next question relates to the first question but is not necessarily dependent on it. It’s the question of individualization, a topic Lisa also explored in a past blog. Within our Oregon field sites we have documented a variety of gray whale foraging tactics (Torres et al. 2018; Video 1) but we do not know if all gray whales use all the tactics or if different individuals only use certain tactics. While I think it’s unlikely that one whale only uses one tactic all the time, I think we could see an individual use one tactic more often than the others. I reason that there could be two reasons for this pattern. First, it could be a response to resource availability; certain tactics are more efficient than others, this could be because the tactic involves capturing the more nutritious prey or because the behavior is less energetically demanding. Second, foraging tactics are socially learned as calves from their mothers, and hence individuals use those learned tactics more frequently. This pattern of maternally inherited foraging tactics has been documented in other marine mammals (Mann and Sargeant 2009; Estes et al. 2003). These questions between foraging tactic, habitat and individualization also tie into the remaining two questions.

My third question is about the relationship between behavior and body condition. As I’ve discussed in a previous blog, I am interested in assessing the relative energetic costs and benefits of the different foraging tactics. Is one foraging tactic more cost-effective than another (less energy out per energy in)? Ever since our lab’s cetacean behavioral ecology class, I’ve been thinking about how my work relates to niche partitioning theory (Pianka 1974).This theory states that when there is low prey availability, niche partitioning will increase. Niche partitioning can occur across several different dimensions: for instance, prey type, foraging location, and time of day when active. If gray whales partition across the prey type dimension, then different whales would feed on different kinds of prey. If whales partition resources across the foraging location dimension, individuals would feed in different areas. Lastly, if whales partition resources across the time axis, individuals would feed at different times of day. Using different foraging tactics to feed on different prey would be an example of partitioning across the prey type dimension. If there is a more preferable prey type, then maybe in years of high prey availability, we would see most of the gray whales using the same tactics to feed on the same prey type. However, in years of low prey availability we might expect to see a greater variety of foraging tactics being used. The question then becomes, does any whale end up using the less beneficial foraging tactic? If so, which whales use the less beneficial tactic? Do the same individuals always switch to the less beneficial tactic? Is there a common characteristic among the individuals that switched, like sex, age, size, or reproductive status? Lemos et al. (2020) hypothesized that the decline in body condition observed from 2016 to 2017 might be a carryover effect from low prey availability in 2016. Could it be that the whales that use the less beneficial tactic exhibit poor body condition the following year?

My fourth, and final, question asks if foraging tactics are passed down from moms to their calves. We have some footage of a mom foraging with her calf nearby, and occasionally it looks like the calf could be copying its mother. Reviewing this footage spiked my interest in seeing if there are similarities between the behavior tactics used by moms and those used by their calves after they have been weaned. While this question clearly relates to the question of individualization, it is also related to body condition: what if the foraging tactics used by the mom is influenced by her body condition at the time?

I hope to answer some of these fascinating questions using the data we have collected during our long field days over the past 6 years. In all likelihood, the story that comes together during my thesis research will be different from what I envision now and will likely lead to more questions. That being said, I’m excited to see how the story unfolds and I look forward to sharing the evolving ideas and plot lines with all of you.

References

Estes, J A, M L Riedman, M M Staedler, M T Tinker, and B E Lyon. 2003. “Individual Variation in Prey Selection by Sea Otters: Patterns, Causes and Implications.” Source: Journal of Animal Ecology. Vol. 72.

Mann, Janet, and Brooke Sargeant. 2009. “ Like Mother, like Calf: The Ontogeny of Foraging Traditions in Wild Indian Ocean Bottlenose Dolphins ( Tursiops Sp.) .” In The Biology of Traditions, 236–66. Cambridge University Press. https://doi.org/10.1017/cbo9780511584022.010.

Pianka, Eric R. 1974. “Niche Overlap and Diffuse Competition” 71 (5): 2141–45.

Soledade Lemos, Leila, Jonathan D Burnett, Todd E Chandler, James L Sumich, and Leigh G. Torres. 2020. “Intra‐ and Inter‐annual Variation in Gray Whale Body Condition on a Foraging Ground.” Ecosphere 11 (4). https://doi.org/10.1002/ecs2.3094.

Torres, Leigh G., Sharon L. Nieukirk, Leila Lemos, and Todd E. Chandler. 2018. “Drone up! Quantifying Whale Behavior from a New Perspective Improves Observational Capacity.” Frontiers in Marine Science 5 (SEP). https://doi.org/10.3389/fmars.2018.00319.