Category: Gray Whale Individual Behavior and Body Condition

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 Torres, Clara 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 Science, 35(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

What drives individual specialization?

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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References

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

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

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

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[12]     D. E. Schindler, J. R. Hodgson, and J. F. Kitchell, “Density-dependent changes in individual foraging specialization of largemouth bass,” Oecologia, vol. 110, no. 4, pp. 592–600, May 1997, doi: 10.1007/s004420050200.

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[14]     L. Kernaléguen, J. P. Y. Arnould, C. Guinet, and Y. Cherel, “Determinants of individual foraging specialization in large marine vertebrates, the Antarctic and subantarctic fur seals,” J. Anim. Ecol., vol. 84, no. 4, pp. 1081–1091, 2015, doi: 10.1111/1365-2656.12347.

[15]     E. M. Keen and K. M. Qualls, “Respiratory behaviors in sympatric rorqual whales: the influence of prey depth and implications for temporal access to prey,” J. Mammal., vol. 99, no. 1, pp. 27–40, Feb. 2018, doi: 10.1093/jmammal/gyx170.

[16]     R. H. MacArthur and E. R. Pianka, “On Optimal Use of a Patchy Environment,” Am. Nat., vol. 100, no. 916, pp. 603–609, 1966, doi: 10.1086/282454.

[17]     C. Fontaine, C. L. Collin, and I. Dajoz, “Generalist foraging of pollinators: diet expansion at high density,” J. Ecol., vol. 96, no. 5, pp. 1002–1010, 2008, doi: 10.1111/j.1365-2745.2008.01405.x.

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[19]     M. M. Pires, P. R. Guimarães Jr, M. S. Araújo, A. A. Giaretta, J. C. L. Costa, and S. F. dos Reis, “The nested assembly of individual-resource networks,” J. Anim. Ecol., vol. 80, no. 4, pp. 896–903, 2011, doi: 10.1111/j.1365-2656.2011.01818.x.

[20]     M. S. Araújo, D. I. Bolnick, and C. A. Layman, “The ecological causes of individual specialisation,”Ecol. Lett., vol. 14, no. 9, pp. 948–958, 2011, doi: https://doi.org/10.1111/j.1461-0248.2011.01662.x.

[21]     C. E. Sheppard, R. Heaphy, M. A. Cant, and H. H. Marshall, “Individual foraging specialization in group-living species,” Anim. Behav., vol. 182, pp. 285–294, Dec. 2021, doi: 10.1016/j.anbehav.2021.10.011.

[22]     S. Wild, S. J. Allen, M. Krützen, S. L. King, L. Gerber, and W. J. E. Hoppitt, “Multi-network-based diffusion analysis reveals vertical cultural transmission of sponge tool use within dolphin matrilines,” Biol. Lett., vol. 15, no. 7, p. 20190227, Jul. 2019, doi: 10.1098/rsbl.2019.0227.

[23]     L. M. Aplin, D. R. Farine, J. Morand-Ferron, A. Cockburn, A. Thornton, and B. C. Sheldon, “Experimentally induced innovations lead to persistent culture via conformity in wild birds,” Nature, vol. 518, no. 7540, pp. 538–541, Feb. 2015, doi: 10.1038/nature13998.

[24]     E. Van de Waal, C. Borgeaud, and A. Whiten, “Potent Social Learning and Conformity Shape a Wild Primate’s Foraging Decisions,” Science, Apr. 2013, doi: 10.1126/science.1232769.

[25]     D. I. Bolnick et al., “Why intraspecific trait variation matters in community ecology,” Trends Ecol. Evol., vol. 26, no. 4, pp. 183–192, Apr. 2011, doi: 10.1016/j.tree.2011.01.009.

Drones with lasers: almost as cool as “sharks with laser beams attached to their heads”

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

The recent advancement in drones (or unoccupied aircraft systems, UAS) has greatly enhanced opportunities for scientists across a broad range of disciplines to collect high-resolution aerial imagery. Wildlife researchers in particular have utilized this technology to study large elusive animals, such as whales, to observe their behavior (see Clara Bird’s blog) and obtain morphological measurements via photogrammetry (see previous blog for a brief history on photogrammetry and drones). However, obtaining useful measurement data is not as easy as flying the drone and pressing record. For this blog, I will provide a brief overview on the basics of using photogrammetry to extract morphological measurements from images collected with drones, as well as the associated uncertainty from using different drone platforms. 

During my PhD at Duke University, I co-developed an open-source photogrammetry software called MorphoMetriX to measure whales in images I collected using drones (Fig. 1) (Torres and Bierlich, 2020) (see this blog for some fieldwork memoirs!). The software is designed to be flexible, simple to use, and customizable without knowledge of scripting languages. Using MorphoMetriX, measurements are made in pixels and then multiplied by the ground sampling distance (GSD) to convert to standard units (e.g., meters) (Fig. 2A). GSD represents the distance on the ground each pixel represents (i.e., the linear size of the pixel) and therefore sets the scale of the image (i.e., cm per pixel). Figure 2A describes how GSD is dependent on the camera sensor, focal length lens, and altitude. Thus, drones equipped with different cameras and focal length lenses will have inherent differences in GSD as altitude increases (Fig. 2B). A larger GSD increases the length each pixel represents in a photo and results in a lower resolution image, potentially obscuring important features in the photo and introducing measurement error.

Figure 1. An example of a Pacific Coast Feeding Group gray whale measured in MorphoMetriX (Torres & Bierlich, 2020).
Figure 2: Overview of photogrammetry methods and calculating ground sampling distance (GSD). A) Photogrammetry methods for how each image is scaled to convert measurements in pixels to standard units (e.g., meters). Altitude is the distance between the camera lens and whale (usually at the surface of the water). Figure from Torres and Bierlich (2020). B) The exact (not accounting for distortion or altitude error) ground sampling distance (GSD) for two drone platforms commonly used to obtain morphological measurements of cetaceans. The difference in GSD between the P4Pro and Inspire 1 is due to the difference in sensor width and focal lengths of the cameras used. Figure from Bierlich et al. (2021).

Obtaining accurate altitude information is a key component in obtaining accurate measurements. All drones are equipped with a barometer, which measures altitude from changes in pressure. In general, barometers usually yield low accuracy in the altitude recorded, particularly for low-cost sensors commonly found on small, off-the-shelf drones (Wei et al., 2016). Dawson et al. (2017) added a laser altimeter (i.e., LightWare SF11/C, https://www.mouser.com//datasheet//2//321//28054-SF11-Laser-Altimeter-Manual-Rev8-1371857.pdf) to a drone, which yields higher accuracy in the altitude recorded. Since then, several studies have adopted use of a laser altimeter to study different species of baleen whales (i.e., Gough et al., 2019; Christiansen et al., 2018).

The first chapter of my dissertation, which was published last year in Marine Ecology Progress Series, compared the accuracy of several drones equipped with different camera sensors, focal length lenses, and a barometer vs. laser altimeter (Bierlich et al., 2021). We flew each drone over a known sized object floating at the surface and collected images at various altitudes (between 10 – 120 m). We used the known size of the floating object to determine the percent error of each measurement at each altitude. We found that 1) there is a lot of variation in measurement error across the different drones when using a barometer to measure altitude and 2) using a laser altimeter dramatically reduces measurement error for each drone (Fig. 3).

Figure 3. The % error for measurements from different drones. Black dashed line represents 0% error (true length = 1.48 m). The gray dashed lines represent under- and over-estimation of the true length by 5% (1.41 and 1.55 m, respectively).

These findings are important because if a study is analyzing measurements that are from more than one drone, the uncertainty associated with those measurements must be taken into account to know if measurements are reliable and comparable. For instance, let’s say we are comparing the body length of two different populations and found that population A is significantly longer than population B. From looking at Figure 3, that significant difference in length between population A and B could be unreliable as the difference may be due to the bias introduced by the type of drone, camera sensor, focal length lens, and whether a barometer or laser altimeter was used for recording altitude. In other words, without incorporating uncertainty associated with each measurement, how do you trust your measurement? 

Hence, the National Institute of Standards and Technology (NIST) states that a measurement is complete only when accompanied by a quantitative statement of its uncertainty (Taylor & Kuyatt, 1994). In our Bierlich et al. (2021) study, we develop a Bayesian statistical model where we use the measurements of the known-sized object floating at the surface (what was used for Fig. 3) as training data to predict the lengths of unknown-sized whales. This Bayesian approach views data and the underlying parameters that generated the data (such as the mean or standard deviation) as random, and thus can be described by a statistical distribution. Using Bayes’ Theorem, a model of the observed data (called the likelihood function), is combined with prior knowledge pertaining to the underlying parameters (called the prior probability distribution) to form the posterior probability distribution, which serves as updated knowledge about the underlying parameter. For example, if someone told me they saw a 75 ft blue whale, I would not be phased. But if someone told me they saw a 150 ft blue whale, I would be skeptical – I’m using prior knowledge to determine the probability of this statement being true. 

The posterior probability distribution produced by this Bayesian approach can also serve as new prior information for subsequent analyses. Following this framework, we used the known-sized objects to first estimate the posterior probability distribution for error for each drone. We then used that posterior probability distribution for error specific to each drone platform as prior information to form a posterior predictive distribution for length of unknown-sized whales. The length of an individual whale can then be described by the mean of this second posterior predictive distribution, and its uncertainty defined as the variance or an interval around the mean (Fig. 4). 

Figure 4. An example of a posterior predictive distribution for total length of an individual blue whale. 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. 

For over half a decade, the GEMM Lab has been collecting drone images of Pacific Coast Feeding Group (PCFG) of gray whales off the coast of Oregon to measure their morphology and body condition (see GRANITE Project Blog). We have been using several different types of drones equipped with different cameras, focal length lenses, barometers, and laser altimeters. These measurements from different drones will inherently have different levels of error associated with them, so adapting these methods for incorporating uncertainty will be key to ensure our measurements are comparable and analyses are robust. To do this, we fly over a known-sized board (1 m) at the start of each flight to use as training data to generate a posterior predictive distribution for length of the an unknown-sized PCFG gray whale that we fly over (Fig. 5). Likewise, we are working closely with several other collaborators who are also using different drones. Incorporating measurement uncertainty from drones used across research labs and in different environments will help ensure robust analyses and provide great opportunity for some interesting comparisons – such as differences in gray whale body condition on their feeding grounds in Oregon vs. their breeding grounds in Baja, Mexico, and morphological comparisons with other baleen whale species, such as blue and humpback whales. We are currently wrapping up measurement from thousands of boards (Fig. 5) and whales (Fig. 1) from 2016 – 2021, so stay tuned for the results!

Figure 5. An example of a known-sized object (1 m long board) used as training data to assess measurement uncertainty. 

References

Bierlich, K.C., Schick, R.S., Hewitt, J., Dale, J., Goldbogen, J.A., Friedlaender, A.S., Johnston D.J. (2021). A Bayesian approach for predicting photogrammetric uncertainty in morphometric measurements derived from UAS. Marine Ecology Progress Series. DOI: https://doi.org/10.3354/meps13814

Christiansen F, Vivier F, Charlton C, Ward R, Amerson A, Burnell S, Bejder L (2018) Maternal body size and condition determine calf growth rates in southern right whales. Mar Ecol Prog Ser 592: 267−281

Dawson SM, Bowman MH, Leunissen E, Sirguey P (2017) Inexpensive aerial photogrammetry for studies of whales and large marine animals. Front Mar Sci 4: 366

Gough, W.T., Segre, P.S., Bierlich, K.C., Cade, D.E., Potvin, J., Fish, F. E., Dale, J., di Clemente, J., Friedlaender, A.S., Johnston, D.W., Kahane-Rapport, S.R., Kennedy, J., Long, J.H., Oudejans, M., Penry, G., Savoca, M.S., Simon, M., Videsen, S.K.A., Visser, F., Wiley, D.N., Goldbogen, J.A. (2019). Scaling of swimming performance in baleen whales. Journal of Experimental Biology, 222(20).https://doi.org/10.1242/jeb.204172  

Taylor, B. N., and Kuyatt, C. E. (1994). Guidelines for Evaluating and Expressing the Uncertainty of NIST Measurement Results. Washington, DC: National Institute of Standards and Technology. 1–25.

Torres, W.I., & Bierlich, K.C. (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  

Wei S, Dan G, Chen H (2016) Altitude data fusion utilizing differential measurement and complementary filter. IET Sci Meas Technol (Singap) 10: 874−879

The costs and benefits of automated behavior classification

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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References

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Figure 2. An asp viper

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

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

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

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References

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

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

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

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

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

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

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

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

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

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

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

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

Blue whales offshore of Monterey, California. 

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

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

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

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

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

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

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

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

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

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

Gray whales off Newport, Oregon

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

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

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

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

References: 

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Little whale, big whale, swimming in the water: A quick history on how aerial photogrammetry has revolutionized the ability to obtain non-invasive measurements of whales

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

The morphology and body size of an animal is one of the most fundamental factors for understanding a species ecology. For instance, fish body size and fin shape can influence its habitat use, foraging behavior, prey type, physiological performance, and predator avoidance strategies (Fig 1). Morphology and body size can thus reflect details of an individual’s current health, likelihood of survival, and potential reproductive success, which directly influences a species life history patterns, such as reproductive status, growth rate, and energetic requirements. Collecting accurate morphological measurements of individuals is often essential for monitoring populations, and recent studies have demonstrated how animal morphology has profound implications for conservation and management decisions, especially for populations inhabiting anthropogenically-altered environments (Miles 2020) (Fig. 1). For example, in a study on the critically endangered European eel, De Meyer et al. (2020) found that different skulls sizes were associated with different ecomorphs (a local variety of a species whose appearance is determined by its ecological environment), which predicted different diet types and resulted with some ecomorphs having a greater exposure to pollutants and toxins than others. However, obtaining manual measurements of wild animal populations can be logistically challenging, limited by accessibility, cost, danger, and animal disturbance. These challenges are especially true for large elusive animals, such as whales that are often in remote locations, spend little time at the surface of the water, and their large size can preclude safe capture and live handling.

Figure 1. Top) A pathway framework depicting how the morphology of an animal influences its habitat use, behavior, foraging, physiology, and performance. These traits all affect how successful an animal is in its environment and can reflect an individual’s current health, likelihood of survival, and potential reproductive success. This individual success can then be scaled up to assess overall population health, which in turn can have direct implications for conservation. Bottom) an example of morphological differences in fish body size and fin shape from Walker et al. (2013). Fineness ratio (f) = length of body ­÷ max body width. 

Photogrammetry is a non-invasive method for obtaining accurate morphological measurements of animals from photographs. The two main types of photogrammetry methods used in wildlife biology are 1) single camera photogrammetry, where a known scale factor is applied to a single image to measure 2D distances and angles and 2) stereo-photogrammetry, where two or more images (from a single or multiple cameras) are used to recreate 3D models. These techniques have been used on domestic animals to measure body condition and estimate weight of dairy cows and lactating Mediterranean buffaloes (Negretti et al., 2008; Gaudioso et al., 2014) and on wild animals to measure sexual dimorphism in Western gorillas (Breuer et al., 2007), shoulder heights of elephants (Schrader et al., 2006), nutritional status of Japanese macaques (Kurita et al., 2012), and the body condition of brown bears (Shirane et al., 2020). Over 70 years ago, Leedy (1948) encouraged wildlife biologists to use aerial photogrammetry from aircraft for censusing wild animal populations and their habitats, where photographs can be collected at nadir (straight down) or an oblique angle, and the scale can be calculated by dividing the focal length of the camera by the altitude or by using a ratio of selected points in an image of a known size. Indeed, aerial photogrammetry has been wildly adopted by wildlife biologists and has proven useful in obtaining measurements in large vertebrates, such as elephants and whales.

Whitehead & Payne (1978) first demonstrated the utility of using aerial photogrammetry from airplanes and helicopters as a non-invasive technique for estimating the body length of southern right whales. Prior to this technique, measurements of whales were traditionally limited to assessing carcasses collected from scientific whaling operations, or opportunistically from commercial whaling, subsistence hunting, stranding events, and bycatch. Importantly, aerial photogrammetry provides a method to collect measurements of whales without killing them. This approach has been widely adopted to obtain body length measurements on a variety of whale and dolphin species, including bowhead whales (Cubbage & Calambokidis, 1987), southern right whales (Best & Rüther, 1992), fin whales (Ratnaswamy and Wynn, 1993), common dolphins (Perryman and Lynn, 1993), spinner dolphins (Perryman & Westlake 1998), and killer whales (Fearnbach et al. 2012). Aerial photogrammetry has also been used to measure body widths to estimate nutritive condition related to reproduction in gray whales (Perryman and Lynn, 2002) and northern and southern right whales (Miller et al., 2012). However, these studies collected photographs from airplanes and helicopters, which can be costly, limited by weather and infrastructure to support aircraft research efforts and, importantly, presents a potential risk to wildlife biologists (Sasse 2003). 

The recent advancement and commercialization of unoccupied aircraft systems (UAS, or drones) has revolutionized the ability to obtain morphological measurements from high resolution aerial photogrammetry across a variety of ecosystems (Fig. 2). Drones ultimately bring five transformative qualities to conservation science compared to airplanes and helicopters: affordability, immediacy, quality, efficiency, and safety of data collection. Durban et al. (2015) first demonstrated the utility of using drones for non-invasively obtaining morphological measurements of killer whales in remote environments. Since then, drone-based morphological measurements have been applied to a wide range of studies that have increased our understanding on different whale populations. For example, Leslie et al. (2020) used drone-based measurements of the skull to distinguish a unique sub-species of blue whales off the coast of Chile. Groskreutz et al. (2019) demonstrated how long-term nutritional stress has limited body growth in northern resident killer whales, while Stewart et al. (2021) found a decrease in body length of North Atlantic Right whales since 1981 that was associated with entanglements from fishing gear and may be a contributing factor to the decrease in reproductive success for this endangered population. 

Drone imagery is commonly used to estimate the body condition of baleen whales by measuring the body length and width of individuals. Recently, the GEMM Lab used body length and width measurements to quantify intra- and inter-seasonal changes in body condition across individual gray whales (Lemos et al., 2020). Drones have also been used to measure body condition loss in humpback whales during the breeding season (Christiansen et al., 2016) and to compare the healthy southern right whales to the skinnier, endangered North Atlantic right whales (Christiansen et al., 2020). Drone-based assessments of body condition have even been used to measure how calf growth rate is directly related to maternal loss during suckling (Christiansen et al., 2018), and even estimate body mass (Christiansen et al., 2019). 

Drone-based morphological measurements can also be combined with whale-borne inertial sensing tag data to study the functional morphology across several different baleen whale species. Kahane-Rapport et al. (2020) used drone measurements of tagged whales to analyze the biomechanics of how larger whales require longer times for filtering the water through their baleen when feeding. Gough et al. (2019) used size measurements from drones and swimming speeds from tags to determine that a whale’s “walking speed” is 2 meters per second – whether the largest of the whales, a blue whale, or the smallest of the baleen whales, an Antarctic minke whale. Size measurements and tag data were combined by Segre et al. (2019) to quantify the energetic costs of different sized whales when breaching. 

Taken together, drones have revolutionized our ability to obtain morphological measurements of whales, greatly increasing our capacity to better understand how these animals function and perform in their environments. These advancements in marine science are particularly important as these methods provide greater opportunity to monitor the health of populations, especially as they face increased threats from anthropogenic stressors (such as vessel traffic, ocean noise, pollution, fishing entanglement, etc.) and climate change. 

Drone-based photogrammetry is one of the main focuses of the GEMM Lab’s project on Gray whale Response to Ambient Noise Informed by Technology and Ecology (GRANITE). This summer we have been collecting drone videos to measure the body condition of gray whales feeding off the coast of Newport, Oregon (Fig. 2). As we try to understand the physiological stress response of gray whales to noise and other potential stressors, we have to account for the impacts of overall nutritional state of each individual whale’s physiology, which we infer from these body condition estimates. 

Figure 2. Drones can help collect images of whales to obtain morphological measurements using photogrammetry and help us fill knowledge gaps for how these animals interact in their environment and to assess their current health. Bottom photo is an image collected by the GEMM Lab of a gray whale being measured in MorphoMetriX software to estimate its body condition. 

References

Best, P. B., & Rüther, H. (1992). Aerial photogrammetry of southern right whales, Eubalaena australis. Journal of Zoology, 228(4), 595-614.

Breuer, T., Robbins, M. M., & Boesch, C. (2007). Using photogrammetry and color scoring to assess sexual dimorphism in wild western gorillas (Gorilla gorilla). American Journal of Physical Anthropology, 134(3), 369–382. https://doi.org/10.1002/ajpa.20678 

Christiansen, F., Vivier, F., Charlton, C., Ward, R., Amerson, A., Burnell, S., & Bejder, L. (2018). Maternal body size and condition determine calf growth rates in southern right whales. Marine Ecology Progress Series, 592, 267–281. 

Christiansen, F. (2020). A population comparison of right whale body condition reveals poor state of North Atlantic right whale, 1–43. 

Christiansen, F., Dujon, A. M., Sprogis, K. R., Arnould, J. P. Y., & Bejder, L. (2016). Noninvasive unmanned aerial vehicle provides estimates of the energetic cost of reproduction in humpback whales. Ecosphere, 7(10), e01468–18. 

Christiansen, F., Sironi, M., Moore, M. J., Di Martino, M., Ricciardi, M., Warick, H. A., … Uhart, M. M. (2019). Estimating body mass of free-living whales using aerial photogrammetry and 3D volumetrics. Methods in Ecology and Evolution, 10(12), 2034–2044. 

Cubbage, J. C., & Calambokidis, J. (1987). Size-class segregation of bowhead whales discerned through aerial stereo-photogrammetry. Marine Mammal Science, 3(2), 179–185. 

De Meyer, J., Verhelst, P., & Adriaens, D. (2020). Saving the European Eel: How Morphological Research Can Help in Effective Conservation Management. Integrative and Comparative Biology, 23, 347–349. 

Gaudioso, V., Sanz-Ablanedo, E., Lomillos, J. M., Alonso, M. E., Javares-Morillo, L., & Rodr\’\iguez, P. (2014). “Photozoometer”: A new photogrammetric system for obtaining morphometric measurements of elusive animals, 1–10.

Gough, W. T., Segre, P. S., Bierlich, K. C., Cade, D. E., Potvin, J., Fish, F. E., … Goldbogen, J. A. (2019). Scaling of swimming performance in baleen whales. Journal of Experimental Biology, 222(20), jeb204172–11. 

Groskreutz, M. J., Durban, J. W., Fearnbach, H., Barrett-Lennard, L. G., Towers, J. R., & Ford, J. K. B. (2019). Decadal changes in adult size of salmon-eating killer whales in the eastern North Pacific. Endangered Species Research, 40, 1 

Kahane-Rapport, S. R., Savoca, M. S., Cade, D. E., Segre, P. S., Bierlich, K. C., Calambokidis, J., … Goldbogen, J. A. (2020). Lunge filter feeding biomechanics constrain rorqual foraging ecology across scale. Journal of Experimental Biology, 223(20), jeb224196–8. 

Leedy, D. L. (1948). Aerial Photographs, Their Interpretation and Suggested Uses in Wildlife Management. The Journal of Wildlife Management, 12(2), 191. 

Lemos, L. S., Burnett, J. D., Chandler, T. E., Sumich, J. L., and Torres, L. G. (2020). Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11.

Leslie, M. S., Perkins-Taylor, C. M., Durban, J. W., Moore, M. J., Miller, C. A., Chanarat, P., … Apprill, A. (2020). Body size data collected non-invasively from drone images indicate a morphologically distinct Chilean blue whale (Balaenoptera musculus) taxon. Endangered Species Research, 43, 291–304. 

Miles, D. B. (2020). Can Morphology Predict the Conservation Status of Iguanian Lizards? Integrative and Comparative Biology. 

Miller, C. A., Best, P. B., Perryman, W. L., Baumgartner, M. F., & Moore, M. J. (2012). Body shape changes associated with reproductive status, nutritive condition and growth in right whales Eubalaena glacialis and E. australis. Marine Ecology Progress Series, 459, 135–156. 

Negretti, P., Bianconi, G., Bartocci, S., Terramoccia, S., & Verna, M. (2008). Determination of live weight and body condition score in lactating Mediterranean buffalo by Visual Image Analysis. Livestock Science, 113(1), 1–7. https://doi.org/10.1016/j.livsci.2007.05.018 

Ratnaswamy, M. J., & Winn, H. E. (1993). Photogrammetric Estimates of Allometry and Calf Production in Fin Whales, \emph{Balaenoptera physalus}. American Society of Mammalogists, 74, 323–330. 

Perryman, W. L., & Lynn, M. S. (1993). Idendification of geographic forms of common dolphin(\emph{Delphinus Delphis}) from aerial photogrammetry. Marine Mammal Science, 9(2), 119–137. 

Perryman, W. L., & Lynn, M. S. (2002). Evaluation of nutritive condition and reproductive status of migrating gray whales (\emph{Eschrichtius robustus}) based on analysisof photogrammetric data. Journal Cetacean Research and Management, 4(2), 155–164. 

Perryman, W. L., & Westlake, R. L. (1998). A new geographic form of the spinner dolphin, stenella longirostris, detected with aerial photogrammetry. Marine Mammal Science, 14(1), 38–50. 

Sasse, B. (2003). Job-Related Mortality of Wildlife Workers in the United States, 1937- 2000, 1015–1020. 

Segre, P. S., Potvin, J., Cade, D. E., Calambokidis, J., Di Clemente, J., Fish, F. E., … & Goldbogen, J. A. (2020). Energetic and physical limitations on the breaching performance of large whales. Elife, 9, e51760.

Shirane, Y., Mori, F., Yamanaka, M., Nakanishi, M., Ishinazaka, T., Mano, T., … Shimozuru, M. (2020). Development of a noninvasive photograph-based method for the evaluation of body condition in free-ranging brown bears. PeerJ, 8, e9982. https://doi.org/10.7717/peerj.9982 

Shrader, A. M., M, F. S., & Van Aarde, R. J. (2006). Digital photogrammetry and laser rangefinder techniques to measure African elephants, 1–7. 

Stewart, J. D., Durban, J. W., Knowlton, A. R., Lynn, M. S., Fearnbach, H., Barbaro, J., … & Moore, M. J. (2021). Decreasing body lengths in North Atlantic right whales. Current Biology.

Walker, J. A., Alfaro, M. E., Noble, M. M., & Fulton, C. J. (2013). Body fineness ratio as a predictor of maximum prolonged-swimming speed in coral reef fishes. PloS one, 8(10), e75422.

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!

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

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

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

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

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

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

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

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

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

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

References

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

Torres, L. G., & Read, A. J. (2009). Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins ( Tursiops truncatus ) in Florida Bay, Florida. Marine Mammal Science, 25(4), 797–815. https://doi.org/10.1111/j.1748-7692.2009.00297.x