Food for thought: conscious reasoning among foraging gray whales

By Nadia Leal, GEMM Lab summer intern, OSU senior, Department of Fisheries, Wildlife, and Conservation Sciences

The OSU GEMM Lab gray whale foraging ecology project in Port Orford is in its seventh year of research. I have the honor to serve as a field assistant for the project as part of Team “Heck Yeah” for the summer 2021 field season. In doing so, I have been presented with the opportunity to take part in its enduring legacy. It is a legacy characterized by novel discovery, distinguished leadership, and endless adventure. These particular aspects motivated me to pursue this internship. Further, the desire to seek out gray whales (Eschrichtius robustus) — a species epitomizing the ability to exhibit resilience in the face of adversity after having experienced two unusual mortality events (UME) in the past two decades and having recovered from historically low population abundances due to whaling — sparked immeasurable excitement.

Figure 1. Nadia operating the theodolite to calculate the location of a gray whale. Source: A. Dawn.

The skills we are acquiring during this field season are essential to master so that I can pursue my aspirations of becoming a marine conservation biologist. For example, we have learned how to operate a theodolite, which is a surveying tool used regularly in marine mammal research to accurately calculate the location of cetaceans and track their movements (Figure 1). We are also learning how to operate a number of other research equipment, to navigate a tandem kayak using a GPS, to process various forms of data, and to identify gray whales! I have especially enjoyed collecting prey samples and navigating our tandem kayak, as kayaking is a summer tradition for my family and the opportunity to kayak in this context is certainly the high point of this internship. The kayak is named “Robustus” after the scientific name of the gray whale: Eschrichtius robustus! (Figure 2). 

Figure 2. Nadia navigating Robustus, the research kayak.

The Port Orford project aims to determine how gray whale foraging is affected by prey quantity and quality. In fact, gray whales exhibit specificity in their selection of prey on the basis of caloric content (Hildebrand 2020). I am particularly interested in the underlying implications these findings imply: the notion of conscious reasoning and decision-making by individual whales as they seek the most suitable prey for its dietary needs among other options to maximize its survivability. Are gray whales in possession of an awareness that allows them to exhibit intentional preference? Can the behavior be attributed to instinct and/or learned behavior, or to cognition comparable to human preference? These and similar questions are my motivation for studying the realm of marine mammal biology. These questions concern intelligence and evolution, which can be effectively investigated through an analysis of cetacean brain structure, as it likely has compelling relationships to their extensive behavioral abilities (Hof and Van Der Gucht 2007). 

For instance, the brain of the gray whale has expanded and developed extensively over evolutionary time in response to distinct selection pressures. Evidence affirms that the behavioral challenges associated with foraging exert strong selection pressures on the evolution of their brain size and structure (Muller and Montgomery 2019)! Selection pressures associated with social cognition are also believed to have contributed to such growth (Connor et al. 1998; Marino 2002; Shultz and Dunbar 2010 ). Further, their neural organization has increased in complexity, leading to greater function and usage of the cortical portion of the brain, which is the portion responsible for higher level activity (Oelschläger and Oelschläger 2002). 

Figure 3. Structure of humpback whale brain representative of baleen species used to infer about gray whales (Hof and Van Der Gucht 2007). 

Though research about baleen whale brain morphology is not as pervasive as that of toothed whales (due to increased susceptibility of toothed whales to captivity given the feasibility of their capture and subsequent analysis in lab/controlled setting), studies have indicated that the brain of baleen whales share similarities to those of humans (Wade et. al 2012). In particular, similarities exist in the frontal lobe of the brain, which is responsible for the complex activities of self-awareness, reasoning, and behavior, as well as for problem-solving and motivation (Hof and Van Der Gucht 2007) (Figure 3). These findings indicate that baleen whales, including the gray whale, have the capability to exhibit intentional preference and take part in conscious decision-making in the recognition of different prey species. The mechanisms responsible for how gray whales may discern prey likely involve a number of the sensory systems, differing in respect to spatial scale (Torres 2017). Thus, gray whales likely rely on various sensory methods, such as vision, sound perception/reception, chemoreception, or an oceanographic stimulus, at differing scales to locate and discern prey. The sensory method employed is dependent on their distance from prey. 

Though we cannot yet confirm whether and/or how gray whales are capable of distinguishing between prey species, what is certain, is that the gray whale is intelligent and quite similar to us. Moreover, they are representative of strength and endurance, providing lessons we can learn from and qualities we can embody. Despite the threats to the species from fishing gear entanglement, ship collisions, climate change, oil industry developments, and being historically hunted, they have remarkably persisted. Thus, we must ensure the existence of the gray whale so they too may thrive for the rest of time, with healthy lives and habitat that is rightfully theirs.

P.S. I would like to thank the GEMM Lab, Oregon State University, Shalynn Pack, Port Orford Sustainable Seafood, Port Orford Co-op, South Coast Tours, Nicki’s Knick Knacks, Leigh Torres, Lisa Hildebrand, Allison Dawn, Clara Bird, Tom Calvanese, Maddie English, Jasen White, and Damian Amerman-Smith for making the internship as special and memorable as it is/was. 

References

Connor, R. C., Mann, J., Tyack, P. L., and Whitehead, H. (1998). Social evolution in toothed whales. Trends in Ecology and Evolution, 13(6): 228– 232. doi: https://doi.org/10.1016/S0169‐5347(98)01326‐3 

Hildebrand, L. (2020). Tonight’s specials include mysids, amphipods, and more: an examination of the zooplankton prey of Oregon gray whales and its impact on foraging choices and prey selection. Master’s thesis, Oregon State University. 

Hof, P.R., and Van Der Gucht, E. (2007). Structure of the cerebral cortex of the humpback whale, Megaptera novaengliae(Cetacea, Mysticeti, Balaenopteridae). The Anatomical Record 290:1-31 doi: 10.1002/ar.a.20407

Marino, L. (2002). Convergence of complex cognitive abilities in cetaceans and primates. Brain, Behavior, and Evolution59: 21–32. doi:  https://doi. org/10.1159/000063731 

Oelschläger, H.A., and Oelschläger, J.S. (2002). Brains. In: Perrin WF, Wu¨ rsig B, Thewissen JGM, editors. Encyclopedia of marine mammals. San Diego, CA: Academic Press. p 133–158.            

Shultz, S., & Dunbar, R. (2010). Encephalization is not a universal macroevolutionary phenomenon in mammals but is associated with sociality. Proceedings of the National Academy of Sciences of the United States of America 107(50): 21582–21586. doi: https://doi.org/10.1073/ pnas.1005246107 

Torres, L.G. (2017). A sense of scale: foraging cetaceans’ use of scale-dependent multimodal sensory systems. Marine Mammal Science 33: 1170-1193. doi:  10.1111/mms.12426 

Wade, P.R., Reeves, R.R., and Mesnick, S.L. (2012). Social and behavioral factors in cetacean responses to overexploitation: are odontocetes less “resilient” than mysticetes?. The Journal of Marine Biology 2012: 1-15. doi:10.1155/2012/567276

Do gray whales count calories? New GEMM Lab publication compares energetic values of prey available to gray whales on two feeding grounds in the eastern North Pacific

By Lisa Hildebrand, PhD student, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Predators have high energetic requirements that must be met to ensure reproductive success and population viability. For baleen whales, this task is particularly challenging since their foraging seasons are typically limited to short temporal windows during summer months when they migrate to productive high latitude environments. Foraging success is a balancing act whereby baleen whales must maximize the amount of energy they intake, while minimizing the amount of energy they expend to obtain food. Maximization of energy intake can be achieved by targeting the most beneficial prey. How beneficial a particular prey type (or prey patch) is can depend on a number of factors such as abundance, density, quality, size, and availability. Determining why baleen whales target particular prey types or patches is an important factor to enhance our understanding of their ecology and can ultimately aid in informing their management and conservation.

The GEMM Lab has several research projects in Newport and Port Orford, Oregon, on the Pacific Coast Feeding Group (PCFG), which is a sub-group of gray whales from the Eastern North Pacific (ENP) population. While ENP gray whales feed in the Bering, Chukchi, and Beaufort Seas (Arctic) in the summer months, the PCFG utilizes the range from northern California, USA to northern British Columbia, Canada. Our work to date has revealed a number of new findings about the PCFG including that they successfully gain weight during the summer on Oregon foraging grounds (Soledade Lemos et al. 2020). Furthermore, females that consistently use the PCFG range as their foraging grounds have successfully reproduced and given birth to calves (Calambokidis & Perez 2017). Yet, the abundance of the PCFG (~250 individuals; Calambokidis et al. 2017) is two orders of magnitude smaller than the ENP population (~20,000; Stewart & Weller 2021). So, why do more gray whales not use the PCFG range as their foraging grounds when it provides a shorter migration while also allowing whales to meet their high energetic requirements and ensure reproductive success? There are several hypotheses regarding this ecological mystery including that prey abundance, density, quality, and/or availability are higher in the Arctic than in the PCFG range, thus justifying the much larger number of gray whales that migrate further north for the summer feeding season. 

Figure 1. Locations of prey samples collected with a light trap (open circles) or opportunistic collections of surface swarms of crab larvae (black triangles) in Newport, along the Oregon coast in the Pacific Northwest coast of the United States.

Our recent paper in Frontiers in Marine Science addressed the hypothesis that prey quality in the Arctic is higher than that of PCFG prey. To test this hypothesis, we first determined the quality (energetic value) of nearshore Oregon zooplankton species that PCFG gray whales are assumed to feed on (based on observations of fine-scale spatial and temporal overlap of foraging gray whales and sampled zooplankton). We obtained prey samples from nearshore reefs along the Oregon coast (Figure 1) as part of the GRANITE project using a light trap, which is a modified water jug with a weight and two floats attached to it, allowing the trap to sit approximately 1 meter above the seafloor. The trap contained a light which attracted zooplankton and effectively captured epibenthic prey of gray whales. Traps were left to soak overnight in locations where gray whales had been observed feeding extensively and collected the following morning. After identifying each specimen to species level and sorting them into reproductive stages, we used a bomb calorimeter to determine the caloric content of each species by month, year, and reproductive stage. We then compared these values to the literature-derived caloric value of the predominant benthic amphipod species that  ENP gray whales feed on in the Arctic. These comparisons allowed us to extrapolate the caloric values gained from each prey type to estimated energetic requirements of pregnant and lactating female gray whales (Villegas-Amtmann et al. 2017). 

Figure 2. Median caloric content and interquartile ranges by (A) species, (B) reproductive stage, and (C) month. Sizes of the zooplankton images are scaled at actual ratios relative to one another.

So, what did we find? Our sampling along the Oregon coast revealed six predominant zooplankton species: two mysid shrimp (Neomysis rayiiHolmesimysis sculpta), two amphipods (Atylus tridensPolycheria osborni), and two types of crab larvae (Dungeness crab megalopae, porcelain crab larvae). These six Oregon prey species showed significant differences in their caloric values, with N. rayii and Dungeness crab megalopae having significantly higher calories per gram than the other prey species (Figure 2), though Dungeness crab megalopae stood out as the caloric gold mines for feeding gray whales in the PCFG range. Furthermore, month and reproductive stage also influenced  the caloric content of some prey species, with gravid (aka pregnant) female mysid shrimp significantly increasing in calories throughout the summer (Figure 3).

Figure 3. Caloric content of different reproductive stages as a function of day of year (DOY; ranging from June to October) for the mysids Holmesimysis sculpta and Neomysis rayii, and the amphipod Atylus tridens. A. tridens is only represented on one panel due to small sample size of this species for the empty brood pouch and gravid reproductive stages. Asterisks indicate significant regressions (p<0.05).

The comparison of our Oregon prey caloric values to the predominant Arctic amphipod (Ampelisca macrocephala) proved our hypothesis wrong:  Arctic amphipods do not have higher caloric value than Oregon prey, which would have help to explain why many more gray whales feed in the Arctic. We found that two Oregon prey species (N. rayii and Dungeness crab megalopae) have higher caloric values than A. macrocephala. If we translate the caloric contents of these prey to gray whale energetic needs, these differences mean that lactating and pregnant gray whales feeding in the PCFG area would need between 0.7-1.03 and 0.22-0.33 metric tons of prey less per day if they fed on Dungeness crab megalopae or N. rayii, respectively, than a whale feeding on Arctic A. macrocephala (Figure 4). 

Figure 4. Daily prey requirements (A: metric tons; B: number of individuals) needed by pregnant and lactating female gray whales to meet their energetic requirements on the foraging ground. Energetic requirement estimates obtained from Villegas-Amtmann et al. (2017). Note the logarithmic scale of y-axis in panel (B).

If quality were the only prey metric that gray whales used to evaluate which food to eat, then it would make very little sense for so many gray whales to migrate to the Arctic when there are prey types of equal and greater quality available to them in the PCFG range. However, quality is not the only metric that influences gray whale foraging decisions. We therefore posit that the abundance, density, and availability of benthic amphipods in the Arctic are higher than the prey species found in the PCFG range. In fact, knowledge of the pulsed reproductive cycle of Dungeness and porcelain crabs allows us to conclude that the larvae of these two species are only available for a few weeks in the late spring and early summer on the Oregon coast. While mysid shrimp, such as N. rayii, are continuously available in the PCFG range throughout the summer, they may occur in less dense and more patchy aggregations than Arctic benthic amphipods. However, current estimates of prey density and abundance for either region are not available, and we do not have data on the energetic costs of the different foraging strategies. While there are still several unknowns, we have documented that higher prey quality in the Arctic is not the reason for the difference in gray whale foraging ground use in the eastern North Pacific.

References

Calambokidis, J., & Perez, A. 2017. Sightings and follow-up of mothers and calves in the PCFG and implications for internal recruitment. IWC Report SC/A17/GW/04 for the Workshop on the Status of North Pacific Gray Whales (La Jolla: IWC).

Calambokidis, J., Laake, J., & Perez, A. 2017. Updated analysis of abundance and population structure of seasonal gray whales in the Pacific Northwest, 1996-2015. IWC Report SC/A17/GW/05 for the Workshop on the Status of North Pacific Gray Whales (La Jolla: IWC).

Soledade Lemos, L., Burnett, J. D., Chandler, T. E., Sumich, J. L., & Torres, L. G. 2020. Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11(4):e03094.

Stewart, J. D., & Weller, D. W. 2021. Abundance of eastern North Pacific gray whales 2019/2020. Department of Commerce, NOAA Technical Memorandum NMFS-SWFSC-639. United States: NOAA. doi:10.25923/bmam-pe91.

Villegas-Amtmann, S., Schwarz, L. K., Gailey, G., Sychenko, O., & Costa, D. P. 2017. East or west: the energetic cost of being a gray whale and the consequence of losing energy to disturbance. Endangered Species Research 34:167-183.

Rock-solid GRANITE: Scaling the disturbance response of individual whales up to population level impacts

By Lisa Hildebrand, PhD student, OSU Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

Since early May, much of the GEMM Lab has been consumed by the GRANITE project, which stands for Gray whale Response to Ambient Noise Informed by Technology and Ecology. Two weeks ago, PhD student Clara Bird discussed our field work preparations, and since May 20th we have conducted five successful days of field work (and one unsuccessful day due to fog). If you are now expecting a blog about the data we have collected so far and whales we encountered, I am sorry to disappoint you. Rather, I want to take a big step back and provide the context of the GRANITE project as a whole, explain why this project and data collection is so important, and discuss what it is that we hope to achieve with our ever-growing, multidisciplinary dataset and team.

We use the Pacific Coast Feeding Group (PCFG) of gray whales that forage off the Oregon coast as our study system to better understand the ecological and physiological response of baleen whales to multiple stressors. Our field methodology includes replicate physiological and ecological sampling of this accessible baleen whale population with synoptic measurement of multiple types of stressors. We collect fecal samples for hormone analysis, conduct drone overflights of whales to collect body condition and behavioral data, record the ambient soundscape through deployment of two hydrophones, and conduct whale photo-identification to link all data streams to each individual whale of known sex, estimated age, and reproductive status. We resample these data from multiple individuals within and between summer foraging seasons, while exposed to different potential stressors occurring at different intensities and temporal periods and durations. The hydrophones are strategically placed with one in a heavily boat-trafficked (and therefore noisy) area close to the Port of Newport, while the second is located in a relatively calm (and therefore quieter) spot near the Otter Rock Marine Reserve (Fig. 1). These hydrophones provide us with information about both natural (e.g. killer whales, wind, waves) and anthropogenic (e.g. boat traffic, seismic survey, marine construction associated with PacWave wave energy facility development) noise that may affect gray whales. During sightings with whales, we also drop GoPro cameras and sample for prey to better understand the habitats where whales forage and what they might be consuming.

Figure 1. Map of GRANITE study area from Seal Rock to Lincoln City with gray whale sightings (yellow circles) and and fecal samples collected (red triangles) from the 2020 field season. Green stars represent the two hydrophone locations. Source: L. Torres.

GEMM Lab PI Dr. Leigh Torres initiated this research project in 2015 and established partnerships with acoustician Dr. Joe Haxel and (then) PhD student Dr. Leila Lemos. Since then, the team working on this project has grown considerably to provide expertise in the various disciplines that the project integrates. Leigh is currently joined at the GRANITE helm by 4 co-PIs: Dr. Haxel, endocrinologist Dr. Kathleen Hunt, biological statistician Dr. Leslie New, and physiologist Dr. Loren Buck. Drs. Alejandro Fernandez Ajo, KC Bierlich and Enrico Pirotta are postdoctoral scholars who are working on the endocrinology, photogrammetry, and biostatistical modelling components, respectively. Finally, Clara and myself are partially funded through this project for our PhD research, with Clara focusing on the links between behavior, body condition, individualization, and habitat, while I am tackling questions about the recruitment and site fidelity of the PCFG (more about these topics below). 

Faculty Research Assistant Todd Chandler supervises PhD student Clara Bird during her maiden drone flight over a whale. Source: L. Torres.

The ultimate goal of this project is to use the PCFG as a case study to quantify baleen whale physiological response to different stressors and model the subsequent impacts on the population by implementing our long-term, replicate dataset into a framework called Population consequences of disturbance (PCoD; Fig. 2). PCoD is built upon the underlying concept that changes in behavior and/or physiology caused by disturbance (i.e. noise) affect the fitness of individuals by impacting their health and vital rates, such as survival, reproductive success, and growth rate (Pirotta et al. 2018). These impacts at the individual level may (or may not) affect the population as a whole, depending on what proportion of individuals in the population are affected by the disturbance and the intensity of the disturbance effect on each individual. The PCoD framework requires quantification of four stages: a) the physiological and/or behavioral changes that occur as a result of exposure to a stressor (i.e. noise), b) the acute effects of these physiological and/or behavioral responses on individual vital rates, and their chronic effects via individual health, c) the way in which changes in health may affect the vital rates of individuals, and d) how changes in individual vital rates may affect population dynamics (Fig. 2; Pirotta et al. 2018). While four stages may not sound like a lot, the amount and longevity of data needed to quantify each stage is immense. 

Figure 2. Conceptual framework of the population consequences of disturbance (PCoD). Letters (A-D) represent the four stages that require quantification in order for PCoD to be implemented. Each colored box represents external (ecological drivers, stressors) and internal (physiology, health, vital rates, behavior) factors that can change over time that are measured for each individual whale (dashed grey boundary line). The effects are then integrated across all individuals in the population to project their effects on the population’s dynamics. Figure and caption adapted from Pirotta et al. 2018.

The ability to detect a change in behavior or physiology often requires an understanding of what is “normal” for an individual, which we commonly refer to as a baseline. The best way to establish a baseline is to collect comprehensive data over a long time period. With our data collection efforts since 2015 of fecal samples, drone flights and photo identification, we have established useful baselines of behavioral and physiological data for PCFG gray whales. These baselines are particularly impressive since it is typically difficult to collect repeated measurements of hormones and body condition from the same individual baleen whale across multiple years. These repeated measurements are important because, like all mammals, hormones and body condition vary across life history phases (i.e., with pregnancy, injury, or age class) and across time (i.e., good or bad foraging conditions). To achieve these repeated measurements, GRANITE exploits the high degree of intra- and inter-annual site fidelity of the PCFG, their accessibility for study due to their affinity for nearshore habitat use, and the long-term sighting history of many whales that provides sex and approximate age information. Our work to-date has already established a few important baselines. We now know that the body condition of PCFG gray whales increases throughout a foraging season and can fluctuate considerably between years (Soledade Lemos et al. 2020). Furthermore, there are significant differences in body condition by reproductive state, with calves and pregnant females displaying higher body conditions (Soledade Lemos et al. 2020). Our dataset has also allowed us to validate and quantify fecal steroid and thyroid hormone metabolite concentrations, providing us with putative thresholds to identify a stressed vs. not stressed whale based on its hormone levels (Lemos et al. 2020).

PhD student Lisa Hildebrand and GRANITE co-PI Dr. Kathleen Hunt collecting a fecal sample. Source: L. Torres.

We continue to collect data to improve our understanding of baseline PCFG physiology and behavior, and to detect changes in their behavior and physiology due to disturbance events. All these data will be incorporated into a PCoD framework to scale from individual to population level understanding of impacts. However, more data is not the only thing we need to quantify each of the PCoD stages. The implementation of the PCoD framework also depends on understanding several aspects of the PCFG’s population dynamics. Specifically, we need to know whether recruitment to the PCFG population occurs internally (calves born from “PCFG mothers” return to the PCFG) or externally (immigrants from the larger Eastern North Pacific gray whale population joining the PCFG as adults). The degree of internal or external recruitment to the PCFG population should be included in the PCoD model as a parameter, as it will influence how much individual level disturbance effects impact the overall health and viability of the population. Furthermore, knowing residency times and home ranges of whales within the PCFG is essential to understand exposure durations to disturbance events. 

To assess both recruitment and residency patterns of the PCFG, I am undertaking a large photo-identification effort, which includes compiling sightings and photo data across many years, regions, and collaborators. Through this effort we aim to identify calves and their return rate to the population, the rate of new adult recruits to the population, and the spatial residency of individuals in our study system. Although photo-id is a basic, commonplace method in marine mammal science, its role is critical to tracking individuals over time to understand population dynamics (in a non-invasive manner, no less). A large portion of my PhD research will focus on the tedious yet rewarding task of photo-id data management and matching in order to address these pressing knowledge gaps on PCFG population dynamics needed to implement the PCoD model that is an ultimate goal of GRANITE. I am just beginning this journey and have already pinpointed many analytical and logistical hurdles that I need to overcome. I do not anticipate an easy path to addressing these questions, but I am extremely eager to dig into the data, reveal the patterns, and integrate the findings into our rock-solid GRANITE project.  

Funding for the GRANITE project comes from the Office of Naval Research, the Department of Energy, Oregon Sea Grant, the NOAA/NMFS Ocean Acoustics Program, and the OSU Marine Mammal Institute.

References

Lemos, L.S., Olsen, A., Smith, A., Chandler, T.E., Larson, S., Hunt, K., and L.G. Torres. 2020. Assessment of fecal steroid and thyroid hormone metabolites in eastern North Pacific gray whales. Conservation Physiology 8:coaa110.

Pirotta, E., Booth, C.G., Costa, D.P., Fleishman, E., Kraus, S.D., Lusseau, D., Moretti, D., New, L.F., Schick, R.S., Schwarz, L.K., Simmons, S.E., Thomas, L., Tyack, P.L., Weise, M.J., Wells, R.S., and J. Harwood. 2018. Understanding the population consequences of disturbance. Ecology and Evolution 8(19):9934-9946.

Soledade Lemos, L., Burnett, J.D., Chandler, T.E., Sumich, J.L., and L.G. Torres. 2020. Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11(4):e03094.

Wave riders or deep divers: what do cetaceans do in stormy weather?

By Lisa Hildebrand¹ and Samara Haver²

¹PhD Student, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab 

²Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, and Conservation Sciences, Cooperative Institute for Marine Resources Studies, Hatfield Marine Science Center

Many aspects of studying cetacean ecology, behavior, population dynamics, health, and sociality depend on being able to see and/or sample cetaceans when they come to the surface. While this research is not necessarily easy given that cetaceans spend the majority of their time underwater out of human sight, it is definitely feasible, as evidenced by decades of cetacean research. However, in order for researchers to observe cetaceans at the surface they need to get out to sea, and this boat-based effort can realistically only be done in good ocean conditions. Any sea-going individual likely uses the Beaufort sea state (BSS) scale as a measure of ocean conditions. For a full breakdown and excellent explanation of what the BSS is, check out our beloved Alexa’s blog; but for the purposes of this blog all you really need to know is that the smaller the BSS (which starts at 0), the calmer the ocean, and the higher the BSS, the rougher & stormier the ocean. There are two main reasons for conducting cetacean research in low BSS: 1) above a certain threshold (usually BSS 4) it becomes difficult to reliably spot and recognize cetaceans at the surface, thus compromising good data collection, and/or 2) to ensure safety and comfort of the research team. 

So, when the BSS gets too high, us humans usually do not go out to sea to study cetaceans, which means that the cetaceans, for the most part, go unobserved. So, many questions arise about what cetaceans are doing during these rough ocean conditions. What does an increased BSS mean for them? Are they unfazed by big waves and strong winds, or are they affected by the weather and take longer dives or seek out fairer seas? A conversation among friends sparked our curiosity of what cetaceans do in stormy conditions and inspired us to collaborate on this blog. Here, we report on what is and is not known about cetaceans in storms, and discuss some ideas about how best to quantify the effects of rough sea conditions on cetaceans.

Slide the arrows to compare sea conditions (BSS 1 [left] vs BSS 6 [right]) experienced by Alexa, the GEMM Lab marine mammal observer on the May 2019 Northern California Current cruise onboard NOAA ship Bell M. Shimada. Source: A. Kownacki/GEMM Lab.

A literature search of cetaceans during storms did not generate many results, which was not surprising to us given the above reasons about researchers not being able to survey in rough sea conditions. However, we did find a couple of interesting studies about cetacean behavior and distribution after storms.

Changes in foraging behavior

Autumnal storms in Maryland, USA resulted in less frequent and shorter encounters of bottlenose dolphins in the US Mid-Atlantic Bight. However, dolphins spent a significantly higher percentage of their encounters feeding after storms than they did before or during them (Fandel et al. 2020). Similarly, bottlenose dolphins in Mississippi Sound displayed an approximately 15% increase in foraging activity for up to 2 years following Hurricane Katrina (Smith et al. 2013). These changes in foraging behavior are attributed to shifts in distributions and behavior of dolphin prey species as a result of altered environmental conditions (primarily sea surface temperature and salinity) following the hurricanes.

Out-of-habitat events and strandings

An out-of-habitat event occurs when an animal is displaced out of its typical habitat. Seven of these events were reported following Hurricane Rita, which hit the southwest Louisiana coast in 2005, with bottlenose dolphins found in flooded roadside ditches, canals, shallow flooded fields, and a natural creek area (Rosel & Watts 2008). These locations ranged from 2.5 to 11 km inland from the coast of the Gulf of Mexico, where these dolphins were displaced from. It is believed that the animals were carried inland on the storm surge that accompanied Hurricane Rita and were left stranded in areas that held water the longest once it started receding (Rosel & Watts 2008).

One of the roadside ditches where a bottlenose dolphin was trapped in Louisiana following Hurricane Rita. Taken from Rosel & Watts (2008).

There have been two mass strandings of pygmy killer whales that are believed to have been a result of hurricanes. In 1995, five pygmy killer whales stranded (three of which died, while two were successfully refloated) in the British Virgin Islands a day after Hurricane Marilyn (Mignucci-Giannoni et al. 1999). In 2006, six pygmy killer whales (five of which died) stranded in New Caledonia during and after Hurricane Jim (Clua et al. 2014). Both studies hypothesize that increased energetic costs, as a result of attempting to evade the hurricanes, coupled with animals becoming disoriented and ending up in shallow waters, is what caused them to strand. 

While these studies reveal post-storm effects on cetaceans, we still do not know exactly how these individuals behaved during the storms. Did they attempt longer dives to stay away from the rough conditions at the surface, thus becoming disoriented? Or were they behaving normally (i.e. foraging, travelling) and were simply “pushed” into waters that they did not intend to go into? Given that very stormy sea conditions do not allow for visual, boat-based surveys, we need to employ different technologies to study cetacean behavior and distribution during storms.

Passive acoustic monitoring (PAM) is a great tool that can monitor ocean environments for us when the seas are too stormy. Using fixed or mobile platforms, underwater PAM listening devices (hydrophone and data storage) can record sounds in the ocean for us to listen and analyze from shore. With PAM we are able to track the vocalizations of marine mammals as well as other sounds in the environment, such as waves crashing and rain. Anecdotally, we have spent many days at sea in conditions that were too rough for visual observations, but we could safely use our PAM tools to detect cetaceans. So, just because the seas may be too rough to see cetaceans, this fact does not mean that we cannot observe them – we just need to listen instead of look. 

There are many tools that can be used to record underwater sounds, including passive acoustic monitoring (PAM; shown in orange), real-time acoustic data collection (green), and active acoustics (blue.) Source: NOAA Fisheries.

A number of studies have investigated whether whales change their vocalization behavior differently in response to changing ambient sound conditions (for example: Dunlop et al. 2010; Fournet et al. 2018). While research on ocean sound levels is often focused on the impact of human-generated or anthropogenic noise, there are also natural, abiotic sound sources (e.g. wind, rain, ice) that can elevate ambient sound levels. One potential animal response to elevated ambient sound levels is to vocalize at a higher intensity, called the Lombard (or cocktail party) effect. This phenomenon is common for us humans – have you ever been at a party and at some point you realize that you are shouting to someone in order to be heard above the noise of the room? That’s the Lombard effect! Humpback whales in Glacier Bay National Park, Alaska, exhibited the Lombard effect in response to both natural and man-made sounds, but the probability of calling was lower when vessels were present compared to times with only natural sounds (Fournet et al. 2018). It is also possible that whales may vocalize at different frequencies, times, or for shorter durations when the ocean becomes louder, which we can easily track with PAM. Unfortunately, PAM is limited to what we are able to hear, so if we do not hear whales we cannot determine if this result is because their vocalizations are masked by higher intensity sounds, if they stopped vocalizing, or if they left the listening area. 

Animal-borne tags are another kind of autonomous observation tool that could help us understand cetacean behavior and distribution in storms. Admittedly, the logistics of applying tags before an imminent storm are probably complex. However, the development of medium-duration archival tags may provide a good trade-off between deploying tags long enough before a storm begins, thus providing safe working conditions for the research team, while minimizing potential physical impacts to the animals (Szesciorka et al. 2016). There are currently no published tag studies that document cetacean behavior during storms, but a study of a gray-headed albatross, fitted with a satellite transmitter, that successfully foraged during an Antarctic storm (Catry et al. 2004) shows the promise of using animal-borne tags to answer these questions.  

As with many questions about animal behavior, our best option is to combine all of our research tools to piece together evidence about what might be going on in the deep, dark, stormy ocean. Simultaneously collecting acoustic and movement & behavior data through PAM and animal-borne tags, respectively, could allow us to determine how cetaceans behave during storms. While we are probably not poised to tackle these questions right now, perhaps another curious graduate student can take it on for their own PhD research…

References

Catry, P., Phillips, R.A., and J.P. Croxall. Sustained fast travel by a gray-headed albatross (Thalassarchie chrysostoma) riding an Antarctic storm. The Auk 121(4):1208-1213.

Clua, E.E., Manire, C.A., and C. Garrigue. 2014. Biological data of pygmy killer whales (Feresa attenuata) from a mass stranding in New Caledonia (South Pacific) associated with Hurricane Jim in 2006. Aquatic Mammals 40(2):162-172.

Dunlop, R.A., Cato, D.H., and M.J. Noad. 2010. Your attention please: increasing ambient noise levels elicits a change in communication behaviour in humpback whales (Megaptera novaeangliae). Proceedings of the Royal Society B 277(1693):doi.org/10.1098/rspb.2009.2319. 

Fandel, A.D., Garrod, A., Hoover, A.L., Wingfield, J.E., Lyubchich, V., Secor, D.H., Hodge, K.B., Rice, A.N., and H. Bailey. 2020. Effects of intense storm events on dolphin occurrence and foraging behavior. Scientific Reports 10:19247.

Fournet, M.E.H., Matthews, L.P., Gabriele, C.M., Haver, S., Mellinger, D.K., and H. Klinck. 2018. Humpback whales Megaptera novaeangliae alter calling behavior in response to natural sounds and vessel noise. Marine Ecology Progress Series 607:251-268.

Mignucci-Giannoni, A.A., Toyos-González, G. M., Pérez-Padilla, J., Rodríguez-López, M. A., and J. Overing. 1999. Mass stranding of pygmy killer whales (Feresa attenuata) in the British Virgin Islands. Journal of the Marine Biological Association of the United Kingdom 80:759-760.

Rosel, P.E., and H. Watts. 2008. Hurricane impacts on bottlenose dolphins in the northern Gulf of Mexico. Gulf of Mexico Science 25:7.

Smith, C.E., Hurley, B.J., Toms, C.N., Mackey, A.D., Solangi, M., and S.A. Kuczaj II. 2013. Hurricane impacts on the foraging patterns of bottlenose dolphins Tursiops truncatus in Mississippi Sound. Marine Ecology Progress Series 487:231-244.

Szesciorka, A.R., Calambokidis, J., and J.T. Harvey. 2016. Testing tag attachments to increase the attachment duration of archival tags on baleen whales. Animal Biotelemetry 4:18.

Love thy mother: maternal care in cetaceans

By Lisa Hildebrand, PhD student, OSU Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

Last week marked the one year anniversary of the pandemic reality we have all been living. It has been an extremely challenging year, with everyone experiencing different kinds of difficulties and hurdles. One challenge that likely unites the majority of us is having to forego seeing our loved ones. For me personally, this is the longest time I have not seen my family (445 days and counting) and I know I am not alone in this situation. My homesickness started a train of thought about cetacean parental care and inspired me to write a blog about this topic. As you can see from the title, this post focuses on maternal care, rather than parental care. This bias isn’t due to my lack of research on this topic or active exclusion, but rather because there are currently no known cetacean species where paternal participation in offspring production and development extends beyond copulation (Rendell et al. 2019). Thus, this blog is all about the role of mothers in the lives of cetacean calves.

Like humans, cetacean mothers invest a lot of energy into their offspring. Most species have a gestation period of 10 or more months (Oftedal 1997). For baleen whale females in particular, pregnancy is not an easy feat given that they only feed during summer feeding seasons. They must therefore acquire all of the energy they will need for two migrations, birth, and (almost) complete lactation, before they will have access to food on feeding grounds again. For pregnant gray whales, a mere 4% loss in average energy intake on the foraging grounds will prevent her from successfully producing and/or weaning a calf (Villegas-Amtmann et al. 2015), demonstrating how crucial the foraging season is for a pregnant baleen whale. Once a calf is born, lactation ensues, ranging in length between approximately 6-8 months for most baleen whale species to upwards of one or two years in odontocetes (Oftedal 1997). The very short lactation period in baleen whales is offset by the large volume (for blue whales, up to 220 kg per day) and high fat percentage (30-50%) of milk that mothers provide for their calves (Oftedal 1997). In contrast, odontocetes (or toothed whales) have a more prolonged period of lactation with less fatty milk (10-30%). This discrepancy in lactation period lengths is in part because odontocete species do not undertake long migrations, which allows females to feed year-round and therefore allocate energy to nursing young for a longer time. 

Blue whale calf nursing in New Zealand in 2016. Footage captured via unmanned aerial system (UAS; drone) piloted by Todd Chandler for GEMM Lab’s OBSIDIAN project. Source: GEMM Lab.

Aside from the energetically costly task of lactation, cetacean mothers must also assist their calves as they learn to swim. Echelon swimming is a common position of mother-calf pairs whereby the calf is in very close proximity to its mother’s mid-lateral flank and provides calves with hydrodynamic benefits. Studies in bottlenose dolphins have shown that swimming in echelon results in a 24% reduction in mean maximum swim speeds and a 13% decrease in distance per stroke (Noren 2008) for mothers, while concurrently increasing average swim speeds and distance per stroke of calves by 28% and 19%, respectively (Noren et al. 2007). While these studies have only been conducted in odontocete species, echelon swimming is also observed in baleen whales (Smultea et al. 2017), indicating that baleen whale females may experience the same reductions in swimming efficiency. Furthermore, mothers will forgo sleep in the first days after birth (killer whales & bottlenose dolphins; Lyamin et al. 2005) and/or shorten their dive foraging times to accommodate calf diving ability (bottlenose dolphins [Miketa et al. 2018] & belugas [Heide-Jørgensen et al. 2001]). Females must endure these losses in foraging opportunities and decreased swimming efficiency when they are at their most nutritionally stressed to ensure the well-being and success of their offspring.

It is at the time of weaning (when a calf becomes independent), that we start to see differences in the maternal role between baleen and toothed whale mothers. Odontocetes have much stronger sociality than baleen whales causing offspring to stay with their mothers for much longer periods. Among the largest toothed whales, such as killer and sperm whales, offspring stay with their mothers in stable matrilineal units for often a lifetime. Among the smaller toothed whales, such as bottlenose dolphins, maternal kin maintain strong bonds in dynamic fission-fusion societies. In contrast, post-weaning maternal care in baleen whales is limited, with the mother-calf pair typically separating soon after the calf is weaned (Rendell et al. 2019). 

Conceptual diagram depicting where baleen (Mysticeti) and toothed (Odontoceti) whales fall on the continuum of low to high social structure and matrilineal kinship structure. The networks at the top depict long-term datasets of photo-identified individuals (red nodes = females, blue nodes = males, yellow nodes = calves) with thickness of connecting lines representing strength of association between individuals. Figure and caption [adapted] from Rendell et al. 2019.

The long-term impact of social bonds in odontocetes is evident through examples of vertically transmitted behaviors (from mother to calf) in a number of species. For example, the use of three unique foraging tactics (sponge carrying, rooster-tail foraging, and mill foraging) by bottlenose dolphin calves in Shark Bay, Australia, was only significantly explained by maternal use of these tactics (Sargeant & Mann 2009). In Brazil, individuals of four bottlenose dolphin populations along the coast cooperatively forage with artisanal fishermen, which involves specialized and coordinated behaviors from both species. This cooperative foraging tactic among dolphins is primarily maintained across generations via social learning from mothers to calves (Simões-Lopeset al. 2016). The risky tactic of intentional stranding by killer whales on beaches to capture elephant seal pups requires a high degree of skill and high parental investment to reduce the associated risk of stranding (Guinet & Bouvier 1995). 

Evidence for vertical transmission of specialized foraging tactics in baleen whales currently does not exist. Bubble-net feeding is a specialized tactic employed by humpback whales in three oceanic regions where multiple individuals work together to herd and trap prey (Wiley et al. 2011). However, it remains unknown whether this behavior is vertically transmitted. Simultaneous video tags from a mother-calf humpback whale pair in the Western Antarctic Peninsula documented synchrony in dives, with the calf’s track lagging behind the mother’s by 4.5 seconds, suggesting that the calf was following its mother (Tyson et al. 2012). Synchronous diving likely allows calves to observe their mothers and practice their diving, and could offer a pathway for them to mimic foraging behaviors and tactics displayed by mothers. 

While there currently may not be evidence for vertical transmission of specialized foraging tactics among the baleen whales, there is documentation of matrilineal fidelity to both foraging (Weinrich 1998, Barendse et al. 2013, Burnham & Duffus 2020) and breeding grounds (Carroll et al. 2015). Matrilineal site fidelity to foraging grounds is not exclusive to baleen whales and has also been documented in a number of odontocete species (Palsbøll et al. 1997, Turgeon et al. 2012). 

In the GEMM Lab, we are interested in exploring the potential long-term bonds, role and impact of Pacific Coast Feeding Group (PCFG) gray whale mothers on their calves. GEMM Lab PhD student Clara Bird is digging into whether specialized foraging tactics, such as bubble blasts and headstands, are passed down from mothers to calves. I hope to assess whether using the PCFG range as a foraging ground (rather than the Arctic region) is a vertically transmitted behavior or whether environmental factors may play a larger role in the recruitment and dynamics of the PCFG. It will take us a while to get to the bottom of these questions, so in the meantime hug your loved ones if it’s safe to do so or, if you’re in my boat, continue to talk to them virtually until it is safe to be reunited.

References

Barendse, J., Best, P. B., Carvalho, I., and C. Pomilla. 2013. Mother knows best: occurrence and associations of resighted humpback whales suggest maternally derived fidelity to a southern hemisphere coastal feeding ground. PloS ONE 8:e81238.

Burnham, R. E., and D. A. Duffus. 2020. Maternal behaviors of gray whales (Eschrichtius robustus) on a summer foraging site. Marine Mammal Science 36:1212-1230.

Carroll, E. L., Baker, C. S., Watson, M., Alderman, R., Bannister, J., Gaggiotti, O. E., Gröcke, D. R., Patenaude, N., and R. Harcourt. 2015. Cultural traditions across a migratory network shape the genetic structure of southern right whales around Australia and New Zealand. Scientific Reports 5:16182.

Guinet, C., and J. Bouvier. 1995. Development of intentional stranding hunting techniques in killer whale (Orcinus orca) calves at Crozet Archipelago. Canadian Journal of Zoology 73:27-33.

Heide-Jørgensen, M. P., Hammeken, N., Dietz, R., Orr, J., and P. R. Richard. 2001. Surfacing times and dive rates for narwhals and belugas. Arctic 54:207-355.

Lyamin, O., Pryaslova, J., Lance, V., and J. Siegel. 2005. Continuous activity in cetaceans after birth. Nature 435:1177.

Miketa, M. L., Patterson, E. M., Krzyszczyk, E., Foroughirad, V., and J. Mann. 2018. Calf age and sex affect maternal diving behavior in Shark Bay bottlenose dolphins. Animal Behavior 137:107-117.

Noren, S. R. 2008. Infant carrying behavior in dolphins: costly parental care in an aquatic environment. Functional Ecology 22:284-288.

Noren, S. R., Biedenbach, F., Redfern, J. V., and E. F. Edwards. 2007. Hitching a ride: the formation locomotion strategy of dolphin calves. Functional Ecology 22:278-283.

Oftedal, O. T. Lactation in whales and dolphins: evidence of divergence between baleen- and toothed-species. Journal of Mammary Gland Biology and Neoplasia 2:205-230.

Palsbøll, P. J., Heide-Jørgensen, M. P., and R. Dietz. 1996. Population structure and seasonal movements of narwhals, Monodon monoceros, determined from mtDNA analysis. Heredity 78:284-292.

Rendell, L., Cantor, M., Gero, S., Whitehead, H., and J. Mann. 2019. Causes and consequences of female centrality in cetacean societies. Philosophical Transactions of the Royal Society B 374:20180066.

Sargeant, B. L., and J. Mann. 2009. Developmental evidence for foraging traditions in wild bottlenose dolphins. Animal Behavior 78:715-721.

Simões-Lopes, P. C., Daura-Jorge, F. G., and M. Cantor. 2016. Clues of cultural transmission in cooperative foraging between artisanal fishermen and bottlenose dolphins, Tursiops truncatus (Cetacea: Delphinidae). Zoologia (Curitiba) 33:e20160107.

Smultea, M. A., Fertl, D., Bacon, C. E., Moore, M. R., James, V. R., and B. Würsig. 2017. Cetacean mother-calf behavior observed from a small aircraft off Southern California. Animal Behavior and Cognition 4:1-23.

Turgeon, J., Duchesne, P., Colbeck, G. J., Postma, L. D., and M. O. Hammill. 2011. Spatiotemporal segregation among summer stocks of beluga (Delphinapterus leucas) despite nuclear gene flow: implication for the endangered belugas in eastern Hudson Bay (Canada). Conservation Genetics 13:419-433.

Tyson, R. B., Friedlaender, A. S., Ware, C., Stimpert, A. K., and D. P. Nowacek. 2012. Synchronous mother and calf foraging behaviour in humpback whales Megaptera novaeangliae: insights from multi-sensor suction cup tags. Marine Ecology Progress Series 457:209-220.

Villegas-Amtmann, S., Schwarz, L. K., Sumich, J. L., and D. P. Costa. 2015. A bioenergetics model to evaluate demographic consequences of disturbance in marine mammals applied to gray whales. Ecosphere 6:1-19.

Weinrich, M. 1998. Early experience in habitat choice by humpback whales (Megaptera novaeaengliae). Journal of Mammalogy 79:163-170.

Wiley, D., Ware, C., Bocconcelli, A., Cholewiak, D., Friedlaender, A., Thompson, M., and M. Weinrich. 2011. Underwater components of humpback whale bubble-net feeding behavior. Behavior 148:575-602.

Putting Physiological Tools to Work for Whale Conservation

By Alejandro Fernandez Ajo, PhD student at the Department of Biology, Northern Arizona University, Visiting scientist in the GEMM Lab working on the gray whale physiology and ecology project  

About four years ago, I was in Patagonia, Argentina deciding where to focus my research and contribute to whale conservation efforts. At the same time, I was doing fieldwork with the Whale Conservation Institute of Argentina at the “Whale Camp” in Península Valdés. I read tons of papers and talked with my colleagues about different opportunities and gaps in knowledge that I could tackle during my Ph.D. program. One of the questions that caught my attention was about the unknown cause (or causes) for the recurrent high calf mortalities that the Southern Right Whale (SRW) population that breeds at Peninsula Valdés experienced during the 2000s (Rowntree et al. 2013). Still, at that time, I was unsure how to tackle this research question.

Golfo San José, Península Valdés – Argentina. Collecting SRW behavioral data from the cliff’s vantage point. Source: A. Fernandez Ajo.

Between 2003 and 2013, at least 672 SRWs died, of which 91% were calves (Sironi et al. 2014). These mortalities represented an average total whale death per year of 80 individuals in the 2007-2013 period, which vastly exceeded the 8.2 average deaths per year of previous years by a ten-fold increase (i.e., 1993-2002) (Rowntree et al. 2013). In fact, this calf mortality rate was the highest ever documented for any population of large whales. During this period, from 2006 to 2009, I was the Coordinator of the Fauna Area in the Patagonian Coastal Zone Management Plan, and I collaborated with the Southern Right Whale Health Monitoring Program (AKA: The Stranding Program) that conducted field necropsies on stranded whales along the coasts of the Península and collected many different samples including whale baleen.

Southern Right Whale, found stranded in Patagonia Argentina. Source: Instituto de Conservación de Ballenas.

In this process, I learned about the emerging field of Conservation Physiology and the challenges of utilizing traditional approaches to studying physiology in large whales. Basically, the problem is that there is no possible way to obtain blood samples (the gold standard sample type for physiology) from free-swimming whales; whales are just too large! Fortunately, there are currently several alternative approaches for gathering physiological information on large whales using a variety of non-lethal and minimally invasive (or non-invasive) sample matrices, along with utilizing valuable samples recovered at necropsy (Hunt et al. 2013). That is how I learned about Dr. Kathleen Hunt’s novel research studying hormones from whale baleen (Hunt et al., 2018, 2017, 2014). Thus, I contacted Dr. Hunt and started a collaboration to apply these novel methods to understand the case of calf mortalities of the SRW calves in Patagonia utilizing the baleen samples that we recovered with the Stranding program at Península Valdés (see my previous blog post).

What is conservation physiology?

Conservation physiology is a multidisciplinary field of science that utilizes physiological concepts and tools to understand underlying mechanisms of disturbances to solve conservation problems. Conservation physiology approaches can provide sensitive biomarkers of environmental change and allow for targeted conservation strategies. The most common Conservation Physiology applications are monitoring environmental stressors, understanding disease dynamics and reproductive biology, and ultimately reducing human-wildlife conflict, among other applications.

I am now completing the last semester of my Ph.D. program. I have learned much about the amazing field of Conservation Physiology and how much more we need to know to achieve our conservation goals. I am still learning, yet I feel that through my research I have contributed to understanding how different stressors impact the health and wellbeing of whales, and about aspects of their biology that have long been obscured or unknown for these giants. One contribution I am proud of is our recent publication of, “A tale of two whales: putting physiological tools to work for North Atlantic and southern right whales,” which was published in January 2021 as a book chapter in “Conservation Physiology: Applications for Wildlife Conservation and Management” published by Oxford University Press: Oxford, UK.

This book outlines the significant avenues and advances that conservation physiology contributes to the monitoring, management, and restoration of wild animal populations. The book also defines opportunities for further growth in the field and identifies critical areas for future investigation. The text and the contributed chapters illustrate several examples of the different approaches that the conservation physiology toolbox can tackle. In our chapter, “A tale of two whales,” we discuss developments in conservation physiology research of large whales, with the focus on the North Atlantic right whale (Eubalaena glacialis) and southern right whale (Eubalaena australis), two closely related species that differ vastly in population status and conservation pressures. We review the advances in Conservation Physiology that help overcome the challenges of studying large whales via a suite of creative approaches, including photo-identification, visual health assessment, remote methods of assessing body condition, and endocrine research using non-plasma sample types such as feces, respiratory vapor, and baleen. These efforts have illuminated conservation-relevant physiological questions for both species, such as discrimination of acute from chronic stress, identification of likely causes of mortality, and monitoring causes and consequences of body condition and reproduction changes.

Book Overview:

This book provides an overview of the different applications of Conservation Physiology, outlining the significant avenues and advances by which conservation physiology contributes to the monitoring, management, and restoration of wild animal populations. By using a series of global case studies, contributors illustrate how approaches from the conservation physiology toolbox can tackle a diverse range of conservation issues, including monitoring environmental stress, predicting the impact of climate change, understanding disease dynamics, and improving captive breeding, and reducing human-wildlife conflict. The variety of taxa, biological scales, and ecosystems is highlighted to illustrate the far-reaching nature of the discipline and allow readers to appreciate the purpose, value, applicability, and status of the field of conservation physiology. This book is an accessible supplementary textbook suitable for graduate students, researchers, and practitioners in conservation science, ecophysiology, evolutionary and comparative physiology, natural resources management, ecosystem health, veterinary medicine, animal physiology, and ecology.

References

Hunt KE, Fernández Ajó A, Lowe C, Burgess EA, Buck CL. 2021. A tale of two whales: putting physiological tools to work for North Atlantic and southern right whales. In: “Conservation Physiology: Integrating Physiology Into Animal Conservation And Management”, ch. 12. Eds. Madliger CL, Franklin CE, Love OP, Cooke SJ. Oxford University press: Oxford, UK.

Sironi, M., Rowntree, V., Di Martino, M. D., Beltramino, L., Rago, V., Franco, M., and Uhart, M. (2014). Updated information for 2012-2013 on southern right whale mortalities at Península Valdés, Argentina. SC/65b/BRG/06 report presented to the International Whaling Commission Scientific Committee, Portugal. <https://iwc.int/home>.

Rowntree, V.J., Uhart, M.M., Sironi, M., Chirife, A., Di Martino, M., La Sala, L., Musmeci, L., Mohamed, N., Andrejuk, J., McAloose, D., Sala, J., Carribero, A., Rally, H., Franco, M., Adler, F., Brownell, R. Jr, Seger, J., Rowles, T., 2013. Unexplained recurring high mortality of southern right whale Eubalaena australis calves at Península Valdés, Argentina. Marine Ecology Progress Series, 493, 275-289. DOI: 10.3354/meps10506

Hunt KE, Moore MJ, Rolland RM, Kellar NM, Hall AJ, Kershaw J, Raverty SA, Davis CE, Yeates LC, Fauquier DA, et al., 2013. Overcoming the challenges of studying conservation physiology in large whales: a review of available methods. Conserv Physiol 1: cot006–cot006.

Hunt, K.E., Stimmelmayr, R., George, C., Hanns, C., Suydam, R., Brower, H., Rolland, R.M., 2014. Baleen hormones: a novel tool for retrospective assessment of stress and reproduction in bowhead whales (Balaena mysticetus). Conserv. Physiol. 2, cou030. https://doi.org/10.1093/conphys/cou030

Hunt, K.E., Lysiak, N.S., Moore, M.J., Rolland, R.M., 2016. Longitudinal progesterone profiles in baleen from female North Atlantic right whales (Eubalaena glacialis) match known calving history. Conserv. Physiol. 4, cow014. https://doi.org/10.1093/conphys/cow014

Hunt, K.E., Lysiak, N.S., Robbins, J., Moore, M.J., Seton, R.E., Torres, L., Buck, C.L., 2017. Multiple steroid and thyroid hormones detected in baleen from eight whale species. Conserv. Physiol. 5. https://doi.org/10.1093/conphys/cox061

Hunt, K.E., Lysiak, N.S.J., Matthews, C.J.D., Lowe, C., Fernández Ajó, A., Dillon, D., Willing, C., Heide-Jørgensen, M.P., Ferguson, S.H., Moore, M.J., Buck, C.L., 2018. Multi-year patterns in testosterone, cortisol and corticosterone in baleen from adult males of three whale species. Conserv. Physiol. 6, coy049. https://doi.org/10.1093/conphys/coy049

What makes a species, a species?

By Lisa Hildebrand, PhD student, OSU Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

Over the roughly 2.5 years that I have researched the Pacific Coast Feeding Group (PCFG) of gray whales, I have thought more and more about what makes a population, a population. From a management standpoint, the PCFG is currently not considered a separate population or even a sub-population of the Eastern North Pacific (ENP) gray whales. Rather, the PCFG is most commonly referred to as a ‘sub-group’ of the ENP. In my opinion, there are valid arguments both for and against the PCFG being designated as its own population. I will address those arguments briefly at the end of this post, but first, I want you to join on me on a journey that is tangential to my question of ‘what makes a population, a population?’ and one that started at the last Marine Mammal Institute Monthly Meeting (MMIMM).

During 2021’s first MMIMM, our director Dr. Lisa Ballance proposed that we lengthen our monthly meeting duration from 1 hour to 1.5 hours. The additional 30 minutes was to allow for an open-ended, institute-wide discussion of a current hot topic in marine mammal science. This proposal was immediately adopted, and the group dove into a discussion about the discovery of a new baleen whale species in the northeastern Gulf of Mexico: Rice’s whale (Balaenoptera ricei). Let me pause here very briefly to reiterate – the discovery of a new baleen whale species!! The fact that anything as large as 12 m could remain undiscovered in our oceans is really quite fascinating and shows that our scientific quest will likely never run out of discoverable subjects. Anyway, the discovery of this new species is supported by several lines of evidence. Unfortunately (but understandably), MMI’s discussion of these topics had to cease after 30 minutes, however I had more questions. I wanted to know what had sparked the researchers to believe that they had discovered a new cetacean species. 

Scientific illustration of Bryde’s whale. Source: NOAA Fisheries.

I started my research by skimming through some news articles about the Rice’s whale discovery. In a Smithsonian Magazine article, I saw a quote by Dr. Patricia Rosel, the lead author of the study detailing Rice’s whale, that read: “But we didn’t have a skull.”. That quote made me pause. A skull? Is that what it takes to discover and establish a new species? This desired piece of evidence seemed rather puzzling and a little antiquated to me, given that the field of genetics is so advanced now and since it is no longer an accepted practice to kill a wild animal just to study it (i.e., scientific whaling). I backtracked through the article to learn that in the 1990s, renowned marine mammal scientist Dale Rice (after whom Rice’s whale was named) recognized that a small population of baleen whale occurred in the northeastern part of the Gulf of Mexico year-round. At the time, this population was believed to be a sub-population of Bryde’s whale. It wasn’t until 2008, that NOAA scientists were able to conduct a genetic analysis of tissue samples from this population, only to find that these whales were genetically distinct from other Bryde’s whales (Rosel & Wilcox 2014). Yet, this information was not enough for these whales to be established as their own species. A skull really was needed to prove that these whales were in fact a new species. Thankfully (for the scientists) but sadly (for the whale), one of these individuals stranded in Sandy Key, Florida, in 2019, and a dedicated team of stranding responders from Florida Fish and Wildlife, Mote Marine Lab, NOAA, Dolphins Plus, and Marine Animal Rescue Society worked tirelessly in difficult conditions to comprehensively document and preserve this animal. Through the diligent work of this, and previous, stranding response teams, Dr. Rosel and her team were provided the opportunity to examine the skull needed to determine population-status. The science team determined that the bones atop the skull around the blowhole provided evidence that these whales were not only genetically, but also anatomically, different from Bryde’s whales. It was this incident, triggered by that short quote in the Smithsonian article, that brought me to my journey of asking ‘what makes a species, a species?’.

Given that I had just read that Dr. Rosel needed a skull to establish Rice’s whale as its own species, I assumed that my search for ‘how to establish a new species’ would end quickly in me finding a list of requirements, one of which would be ‘must present anatomical/skeletal evidence’. To my surprise, my search did not end quickly, and I did not find a straightforward list of requirements. Instead, I discovered that my question of ‘what makes a species, a species?’ does not have a black-and-white answer and involves a lot of debate.

The skull of this stranded whale was a large piece of evidence in establishing Rice’s whale as its own species. Source: Smithsonian Magazine from NOAA / Florida Fish and Wildlife Commission

Kevin de Queiroz, a vertebrate, evolutionary, and systematic biologist who has published extensively on theoretical and conceptual topics in systematic and evolutionary biology, believes that the issue of species delimitation (‘what makes a species, a species?’) has been made more complicated by a larger issue involving the concept of species itself (‘what is a species?’) (De Queiroz 2007). To date, there are 24 recognized species concepts (Mayden 1997). In other words, there are 24 different definitions of what a species is. Perhaps the most common example is the biological species concept where a species is defined as a group of individuals that are able to produce viable and fertile offspring following natural reproduction. Another example is the ecological species concept whereby a species is a group of organisms adapted to a particular set of resources and conditions, called a niche, in the environment. Problematically, many of these concepts are incompatible with one another, meaning that applying different concepts leads to different conclusions about the boundaries and numbers of species in existence (De Queiroz 2007).

This large number of species concepts is due to the different interests of certain subgroups of biologists. For example, highlighting morphological differences between species is central to paleontologists and taxonomists, whereas ecologists will focus on niche differences. Population geneticists will attribute species differences to genes, while for systematists, monophyly will be paramount. It goes on and on. And so does the debate about the concept of species. It seems that there currently is not one clear, defined consensus on what a species is. Some biologists argue that a species is a species if it is genetically different, while others will insist that skeletal and morphological evidence must be present. From what I can tell, it seems that scientists describe and (attempt to) establish a new species by publishing their lines of evidence, after which experts in the field discuss and evaluate whether a new species should be established. 

In the field of marine mammal science, the Society of Marine Mammalogy’s Taxonomic Committee is charged with maintaining a standard, accepted classification and nomenclature of marine mammals worldwide. The committee annually considers and evaluates new, peer-reviewed literature that proposes changes (including additions) to marine mammal taxonomy. I expect that the case of Rice’s whale will be on the committee’s docket this year. Given that Rosel and co-authors presented geographic, morphological, and genetic evidence to support the establishment of Rice’s whale, I would not be surprised if the committee adds it to their curated list.

After taking this dive into the ‘what makes a species, a species?’ question, let’s see if we can apply some of what we’ve learned to the ‘what makes a population, a population?’ question regarding the PCFG and ENP gray whales. Following the ecological species concept, an argument for the recognition of the PCFG as its own population would be that they occupy an entirely different environment during their summer foraging season than the ENP whales. Not only are the geographic ranges different, but PCFG whales also show behavioral differences in their foraging tactics and targeted prey. The argument against the PCFG being classified as its own population is largely supported by genetic analysis that has revealed ambiguous evidence that the PCFG and ENP are not genetically isolated from one another. While one study has shown that there is maternal cultural affiliation within the PCFG (meaning that calves born to PCFG females tend to return to the PCFG range; Frasier et al. 2011), another has revealed that mixing between ENP and PCFG gray whales on the breeding grounds does occur (Lang et al. 2014). So, even though these two groups feed in areas that are very far apart (ENP: Arctic vs PCFG: US & Canadian west coast) and certain individuals do show a propensity for a specific feeding ground, the genetic evidence suggests that they mix when on their breeding grounds in Baja California, Mexico. Depending on which species concept you align with, you may see better arguments for either side.

PCFG gray whale along the Oregon coast during the GEMM Lab’s 2020 GRANITE summer field season. Image captured under NOAA/NMFS permit #21678. Source: GEMM Lab.

You may be wondering why it is important to even ponder questions like ‘what makes a species, a species?’ and ‘what makes a population, a population?’. Does it really matter if the PCFG are considered their own population? Would anything really change? The answer is, most likely, yes. If the PCFG were to be recognized as their own population, it would likely have an immediate effect on their conservation status and subsequently on how the population needed to be managed. Rather than being under the umbrella of a large, (mostly) stable population of ~25,000 individuals, the PCFG would consist of only ~250 individuals. A group this small would possibly be considered “endangered”, which would require much stricter monitoring and management to ensure that their numbers did not decline from year to year, especially due to anthropogenic activities. 

For a long time, I felt like taxonomy was a bit of an archaic scientific field. In my mind, it was something that biologists had focused their time and energy on in the 18th century (most notably Carl Linnaeus, whose taxonomic classification system is still used today), but something that many biologists have moved on from focusing on in the 21st century. However, as I have developed and grown over the last years as a scientist, I have learned that scientific disciplines are often heavily intertwined and co-dependent on one another. As a result, I am able to see the enormous value and need for taxonomic work as it plays a large part in understanding, managing, and ultimately, conserving species and populations.

Literature cited

De Queiroz, K. 2007. Species concepts and species delimitation. Systematic Biology 56(6):879-886.

Frasier, T. R., Koroscil, S. M., White, B. N., and J. D. Darling. 2011. Assessment of population substructure in relation to summer feeding ground use in the eastern North Pacific gray whale. Endangered Species Research 14:39-48.

Lang, A. R., Calambokidis, J., Scordino, J., Pease, V. L., Klimek, A., Burkanov, V. N., Gearin, P., Litovka, D. I., Robertson, K. M., Mate, B. R., Jacobsen, J. K., and B. L. Taylor. 2014. Assessment of genetic structure among eastern North Pacific gray whales on their feeding grounds. Marine Mammal Science 30(4):1473-1493.

Mayden, R. L. 1997. A hierarchy of species concepts: the denouement of the species problem in The Units of Biodiversity – Species in Practice Special Volume 54 (M. F. Claridge, H. A. Dawah, and M. R. Wilson, eds.). Systematics Association.

Rosel, P. E., and L. A. Wilcox. 2014. Genetic evidence reveals a unique lineage of Bryde’s whale in the northeastern Gulf of Mexico. Endangered Species Research 25:19-34.

GEMM Lab 2020: A Year in the Life

By Lisa Hildebrand, PhD student, OSU Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

Despite the trials and tribulations of 2020, the GEMM Lab has persevered and experienced many successes and high points. Join me, perhaps with a holiday beverage of choice in-hand, for a summary of what the lab and its members have achieved this year.

The GEMM Lab celebrated several milestones this year. We were all extremely excited and proud when halfway through the year, in July, GEMM Lab PI, Dr. Leigh Torres, was promoted to Associate Professor and granted indefinite tenure in the Department of Fisheries & Wildlife. Leigh joined the department in 2014 and has since completed 13 research projects, is leading 10 current research projects, has graduated 7 graduate students, and is currently advising 4 PhD students and a postdoctoral scholar. A big hurrah to Leigh, our inspiring and tireless captain at the GEMM Lab helm!!

Leigh isn’t the only GEMM Lab member to have received a new title. In March, Leila successfully defended her PhD thesis entitled “Body condition and hormone assessment of eastern North pacific gray whales (Eschrichtius robustus) and associations to ambient noise” and thus graduated from being a PhD candidate to being Dr. Leila Soledade Lemos. Leila is currently a postdoctoral associate at Florida International University. I (Lisa Hildebrand) defended my Master’s thesis “Tonight’s specials include mysids, amphipods, and more: An examination of the zooplankton prey of Oregon gray whales and its impact on foraging choices and prey selection” just a few weeks ago and now bear the title of Master of Science. I am excited to announce that I won’t be leaving the GEMM Lab anytime soon as I will continue to  work with Leigh as I pursue my PhD. Our final new title recipient is Dawn who at the start of December advanced to PhD candidacy after successfully passing her written comprehensive exams in mid-November and her oral comprehensive exams in early December.

Summer is a busy time in the GEMM Lab, largely because it is the time when gray whales are distributed along our Oregon coast for their feeding season and therefore when both of our gray whale projects (GRANITE, or Gray whale Response to Ambient Noise Informed by Technology and Ecology, and the Port Orford foraging ecology project) collect another year of data. With the COVID-19 pandemic in its early stages in the spring (when we start to prep for our field seasons), it was uncertain whether we would be able to get into the field at all. However, after weeks of drafting up and submitting COVID-19 safety plans and precautions, Leigh was able to get both of our gray whale field seasons approved to go ahead this summer! This task was not easy since both projects require some form of travel and sampling methods that do not always allow for 6-feet of distance between team members. Furthermore, the Port Orford project requires the whole team to live and work out of OSU’s Port Orford Field Station together. Despite the hurdles, both projects had successful field seasons. If you want to hear more about the specifics of the field seasons, check out the field season summary blog.

Gray whales weren’t the only species to grab our attention in the field this year. OPAL (Overlap Predictions about Large whales) had a successful second year with Leigh and MMI faculty research assistant Craig Hayslip taking to the skies in United States Coast Guard helicopters four times a month. The project seeks to identify co-occurrence between whales and fishing effort in Oregon to reduce entanglement risk. Leigh and Craig documented numerous cetacean species including blue, fin, humpback, sperm whales, and killer whales. To help with this work, we are so excited to officially have Solène Derville back in the GEMM Lab as a postdoctoral scholar who will work on statistical models aimed at predicting habitat use and distribution patterns of whales off the Oregon coast. While our wish to physically welcome Solène back to Oregon this year did not quite pan out, we are hopeful that she will make the journey from New Caledonia to Oregon in 2021!

The data collected during the helicopter flights will be complimented by the marine mammal observer data that various members of the GEMM Lab have collected over the last four years aboard NOAA Ship Bell M. Shimada as part of the Northern California Current Ecosystem survey. These surveys typically occur three times a year (February, May, September). Although the pandemic threw a wrench into the May cruise, the September cruise was able to go ahead with Dawn and Clara on-board as the two marine mammal observers. It was a very successful cruise, with abundant marine mammal sightings and good survey conditions. Read more about those cruises in Clara and Dawn’s blogs.

While the GEMM Lab did not undertake any field work in New Zealand this year, Leigh and Dawn did travel there in February to meet with scientific colleagues, representatives of the oil and gas industry, and environmental managers, including the New Zealand Minister of Conservation, the Honorable Eugenie Sage. The trip allowed Leigh and Dawn to present their research on blue whales and discuss management implications. These meetings have been highly beneficial as they shared their latest research and results to assist with the development of a marine mammal sanctuary within the industrial region where their research is conducted.

The GEMM Lab prides itself on having strong outreach components to our research, ensuring that young students (high school and undergraduate) from diverse backgrounds have an opportunity to learn STEM skills. Some outreach opportunities were not possible in 2020, but the GEMM Lab continued our efforts where possible. Clara taught a photogrammetry workshop for the Marine Studies Initiative student club Ocean11, where students were taught how to measure whales from drone images. The success of the workshop (and earlier iterations of it in 2019) led to Clara turning it into a lab for Dr. Renee Albertson’s FW 469 Physiology/Behavior of Marine Megafauna class. As one of the program coordinators for the Fisheries & Wildlife Mentorship Program, I co-hosted an Intro to R & RStudio workshop this fall. Rachel taught a remote intensive science communication workshop during her first term in grad school. Although COVID-19 meant that one-on-one mentorships had to be a little more distant, over the course of the year, the GEMM Lab still supervised a total of 7 students that assisted our work in a variety of ways (field and/or lab work, data analyses, independent projects) on a number of projects going on in the lab.

In a typical year, GEMM Lab members would have undertaken quite a lot more travel, largely to attend conferences. Due to COVID-19, most conferences were either cancelled or held virtually. Leigh gave the plenary talk at the annual State of the Coast Conference, one of the favorite conferences of the GEMM Lab as it brings together scientists, stakeholders, managers, students, and the public to discuss Oregon-centric topics. Dawn gave an oral presentation at the International Marine Conservation Congress. The talk was titled “Wind, green water, and blue whales: Predictive models forecast blue whale distribution in an upwelling system to mitigate industrial impacts” as part of a symposium focused on evidence-based solutions for the management of large marine vertebrate species. Clara presented at the annual Research Advances in Fisheries, Wildlife & Ecology symposium hosted by the graduate student association in the Department of Fisheries & Wildlife. Clara’s talk, which was about her proposed PhD research, was titled “Drone footage reveals patterns of gray whale behavior across space, time, and the individual”.

While our travel may have been reduced this year, the lab certainly has had a prolific year of writing! The 19 new publications in 16 scientific journals include contributions from Leigh (6), Leila (5), Rachael (4), Solène (3), Clara (3), Dawn (2), and Ale (1). Scroll down to the end of the post to see the full list.

We are also very excited about a new addition to the lab. Rachel Kaplan, who is co-advised by Leigh and Dr. Kim Bernard in the College of Earth, Ocean, and Atmospheric Sciences, started her PhD at OSU in the fall. Rachel is one of this year’s recipients of the highly-competitive National Science Foundation’s Graduate Research Fellowship. Receiving the fellowship allowed Rachel to wrap up her job at the Bigelow Laboratory for Ocean Sciences in Maine and move to Oregon. The journey wasn’t easy (Rachel moved in the midst of the pandemic and during the height of the wildfires that raged across the U.S. West Coast) but she made it here safely! For her PhD, Rachel will try to understand how oceanographic factors and prey patches shape the distribution of whales in Oregon waters (with data collected through the OPAL project) to work towards solutions to the high rates of whale entanglements in fishing gear that have occurred on the West Coast since 2014. Welcome Rachel! 

While we persevered through tough times this year and have been lucky to celebrate many accomplishments, nothing prepared us for the shock that we all felt, and are still feeling deeply, about the loss of our fellow GEMM Lab graduate student Alexa Kownacki just over a month ago. Alexa’s optimism, generosity, and kindness were unparalleled, and the hole that she leaves in the lab and in our lives individually is gaping. The lab wrote a collaborative blog about Alexa a few weeks ago and we have created a website in her honor, where we encourage everyone to post photos, tributes or stories about Alexa. It has been so comforting to us to read people’s memories of Alexa that allow us to learn new things about her and remind us of our own memories. Alexa, we think of you every day and we miss you.

Alexa in her element

If you are reading this post, we would like to say thank you for all the support and interest in our work – we really appreciate it! Our blog’s viewership this year (a whopping 25,588 views!) has increased over a seven-fold since its creation in 2015 (3,462 views). We hope you will continue to join us on our journeys in 2021. Until then, stay safe, mask up & happy holidays from the GEMM Lab!

A GEMM Lab Happy Hour Zoom

Publications

Ajó, A. A. F., Hunt, K. E., Giese, A. C., Sironi, M., Uhart, M., Rowntree, V. J., Marón, C. F., Dillon, D., DiMartino, M., & Buck, C. L. (2020). Retrospective analysis of the lifetime endocrine response of southern right whale calves to gull wounding and harassment: A baleen hormone approach. General and Comparative Endocrinology, 296, 113536.

Albert, C., …, Orben, R. A., et al. (2020). Seasonal variation of mercury contamination in Arctic seabirds: a pan-arctic assessment. Science of the Total Environment, 750, 142201.

Barlow, D. R., Bernard, K. S., Escobar-Flores, P., Palacios, D. M., & Torres, L. G. (2020). Links in the trophic chain: modeling functional relationships between in situ oceanography, krill, and blue whale distribution under different oceanographic regimes. Marine Ecology Progress Series642, 207-225.

Baylis, A. M. M., Tierney, M., Orben, R. A., González de la Peña, D., & Brickle, P. (2020). Non-breeding movements of Gentoo penguins at the Falkland Islands. Ibis, doi:10.1111/ibi.12882.

Bird, C., & Bierlich, K.. (2020).  CollatriX: A GUI to collate MorphoMetriX outputs. Journal of Open Source Software5(51), 2328. doi:10.21105/joss10.21105/joss.02328.

Bird, C., Dawn, A. H., Dale, J., & Johnston, D. W. (2020). A Semi-Automated Method for Estimating Adélie Penguin Colony Abundance from a Fusion of Multispectral and Thermal Imagery Collected with Unoccupied Aircraft Systems. Remote Sensing12(22), 3692. doi:10.3390/rs12223692.

Chero, G., Pradel, R., Derville, S., Bonneville, C., Gimenez, O., & Garrigue, C. (2020). Reproductive capacity of an endangered and recovering population of humpback whales in the Southern Hemisphere. Marine Ecology Progress Series, 643, 219-227.

Derville, S.Torres, L. G., Zerbini, A. N., Oremus, M., & Garrigue, C. (2020). Horizontal and vertical movements of humpback whales inform the use of critical pelagic habitats in the western South Pacific. Scientific Reports, 10, 4871.

DiGiacomo, A. E., Bird, C., Pan, V. G., Dobroski, K., Atkins-Davis, C., Johnston, D. W., & Ridge, J. T.. (2020). Modeling Salt Marsh Vegetation Height Using Unoccupied Aircraft Systems and Structure from Motion. Remote Sensing12(14), 2333. doi:10.3390/rs12142333.

Garrigue, C., Derville, S., Bonneville, C., Baker, C. S., Cheeseman, T., Millet, L., Paton, D., & Steel, D. (2020). Searching for humpback whales in a historical whaling hotspot of the Coral Sea, South Pacific. Endangered Species Research, 42, 67-82.

Hauser-Davis, R. A., Monteiro, F., Chávez da Rocha, R. C., Lemos, L., Duarte Cardoso, M., & Siciliano, S. (2020). Titanium as a contaminant of emerging concern in the aquatic environment and the current knowledge gap regarding seabird contamination. Ornithologia, 11, 7-15.

Hindell, M. A., … Torres, L. G., et al. (2020). Tracking of marine predators to protect Southern Ocean ecosystems. Nature, 580(7801), 87-92.

Jones, K. A., Baylis, A. M. M., Orben, R. A., Ratcliffe, N., Votier, S. C., Newton, J., & Staniland, I. J. (2020). Stable isotope values in South American fur seal pup whiskers as proxies of year-round maternal foraging ecology. Marine Biology, 167(10), 1-11.

Kroeger, C. E., Crocker, D. E., Orben, R. A., Thompson, D. R., Torres, L. G., Sagar, P. M., Sztukowski, L. A., Andriese, T., Costa, D. P., & Shaffer, S. A. (2020). Similar foraging energetics of two sympatric albatrosses despite contrasting life histories and wind-mediated foraging strategies. Journal of Experimental Biology, 223, jeb228585.

Lemos, L. S., Olsen, A., Smith, A., Chandler, T. E., Larson, S., Hunt, K., & Torres, L. G. (2020). Assessment of fecal steroid and thyroid hormone metabolites in eastern North Pacific gray whales. Conservation Physiology, 8, coaa110.

Monteiro, F., Lemos, L. S., et al. (2020). Total and subcellular Ti distribution and detoxification processes in Pontoporia blainvillei and Steno bredanensis dolphins from southeastern Brazil. Marine Pollution Bulletin, 153, 110975.

Quinete, N., Hauser-Davis, R. A., Lemos, L. S., Moura, J. F., Siciliano, S., & Gardinali P. R. (2020). Occurrence and tissue distribution of organochlorinated compounds and polycyclic aromatic hydrocarbons in Magellanic penguins (Spheniscus magellanicus) from the southeastern coast of Brazil. Science of the Total Environment, 749, 141473.

Soledade Lemos, L., Burnett, J. D., Chandler, T. E., Sumich, J. L., & Torres, L. G. (2020). Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere, 11(4), e03094.

Torres, L. G., Barlow, D. R.Chandler, T. E., & Burnett, J. D. (2020). Insight into the kinematics of blue whale surface foraging through drone observations and prey data. PeerJ8, e8906.

Summaries, highlights, and musings – our 2020 gray whale field seasons at a glance

By Lisa Hildebrand, MSc student, OSU Department of Fisheries & Wildlife, Marine Mammal Institute, Geospatial Ecology of Marine Megafauna Lab

Fall has arrived in the Pacific Northwest. For humans, it means packing away the shorts and sandals, and getting the boots, raincoats and firewood ready. For gray whales, it means gulping down the last meal of zooplankton they will eat for several months and commencing the journey to warmer waters and sunnier skies in Mexico where they will spend the winter fasting, calving, and nursing. While the GEMM Lab may still squeeze in a day or two of field work this week, we are slowly wrapping up the 2020 field season as conditions get rougher and our beloved gray whales gradually depart our waters. This year marked the 6th year of data collection for both of our gray whale projects: the Newport project that investigates the impacts of multiple stressors on gray whale ecology and health, and the Port Orford project that explores fine-scale foraging ecology of gray whales and their zooplankton prey. Since it will be several months before the GEMM Lab heads back out onto the water again, I thought I would summarize our two field seasons, share some highlights, and muse about the drivers of our observations this summer.

Summaries

Our RHIB Ruby zipped around the central and southern Oregon coast on 33 different days. The summer started slow, with several days of field work where we encountered no whales despite surveying our entire study region. Our encounters picked up towards the end of June and by the end of the summer we totaled 107 sightings, encountering 46 unique individuals, 36 of which were resightings of known individuals we have identified in previous years. Our Newport star of the summer was Solé, a female gray whale we have seen every year since 2015, and we also saw many of our other regulars including Casper, Rafael, Spray, Bit, and Heart. None of these whales shone as bright as Solé though. We flew the drone over her 8 times and collected 7 fecal samples (one of which was the biggest whale fecal sample I have ever seen!). In total, we collected 30 fecal samples and flew the drone 88 times. These data will allow us to continue measuring body condition and hormone levels of Pacific Coast Feeding Group (PCFG) gray whales that use the Oregon coast.

Our tandem research kayak Robustus may not be as zippy as Ruby (it is powered by human muscle rather than a powerful outboard engine after all), but it certainly continues to be a trusty vessel for the Port Orford team. The Port Orford research team, named the Theyodelers this year, collected 181 zooplankton samples and conducted 180 GoPro drops during the month of August from Robustus. Despite the many samples collected, the size of our prey samples remained relatively small throughout the whole season compared to previous years. The cliff team surveyed for a total of 117 hours, of which 15 were spent tracking whales with the theodolite and resulted in 40 different tracklines of whale movements. The whale situation in Port Orford was similar to the pattern of whale sightings in Newport, with low whale sightings at the start of the field season. Luckily, by the start of August (which marked the start of data collection for the Theyodelers), the number of whales using the Port Orford area, especially the two study sites, Mill Rocks & Tichenor Cove, had increased. Of the whales that came close enough to shore for us to identify using photo-id, we tracked 5 unique individuals, 3 of which we also saw in Newport this year. The Port Orford star of the summer was Smudge, with his tracklines making up a quarter of all of our tracklines collected. Smudge is also the whale we sighted most often last year in Port Orford. 

Highlights

Many of you may be familiar with the whale Scarlett (formally known as Scarback). Scarlett is a female, at least 24 years old (she was first documented  in the PCFG range in 1996), who is well-known (and easily identified) by the large concave injury on her back that is covered in whale lice, or cyamids. No one knows for certain how Scarlett sustained this injury (though there are stories), however what we do know is that it has not prevented this female from reproducing and successfully raising several calves over her lifetime. The GEMM Lab last saw Scarlett with a calf (which we named Brown) in 2016. Since Scarlett is such a famous whale with a unique history, it shouldn’t be a surprise that one of our highlights this summer is the fact that Scarlett showed up with a new calf! In keeping with a “shades of red” theme, Leigh came up with the name Rose for the new calf. In July, the mom-calf pair put on quite a cute performance, with Rose rising up on Scarlett’s back, giving the team a glimpse of its face. The Scarlett-Rose highlight doesn’t end there though. Just last week, we had a very brief encounter in choppy, swelly waters with a small whale. The whale surfaced just twice allowing us to capture photo-id images, and as we were looking around to see where it would come up a third time, it suddenly breached approximately 20 m from the boat. Lo-and-behold, after comparing our photos of the whale to our catalogue, we realized that this elusive, breaching whale was Rose! I am excited to see whether Rose will return to the Oregon coast next summer and become a PCFG regular just like her mom.

The highlight of the field season in Port Orford is the trial, failures and small successes of a new element to the project. There is still a lot that we do not know and understand about PCFG gray whales. One such thing is the way in which gray whales maneuver their large bodies in shallow rocky habitats, often riddled with kelp, and how exactly they capture their zooplankton prey in these environments. Using drones has certainly helped bring some light into this darkness and has led to the documentation of many novel foraging behaviors (Torres et al. 2018). However, the view from above is unable to provide the fine-scale interactions between whales, kelp, reefs, and zooplankton. Instead, we must somehow find a way to watch the whales underwater. Enter CamDo. CamDo is a technology company that designs specialty products to allow for GoPro cameras to be used for time-lapsed recordings over long periods of time in harsh environmental conditions. One of their products is a housing specifically designed for long-term filming underwater – exactly what we need! The journey was not as easy as simply purchasing the housing. We also needed to build a lander for the housing to sit on (thankfully our very own Todd Chandler designed and built something for us), and coordinate with divers and a vessel to deploy and retrieve the set-up, as well as undertake weekly battery and SD cards swaps (thankfully Dave Lacey of South Coast Tours and a very generous group of divers* donated their time and resources to make this happen). We unfortunately had some technological difficulties and bad visibility for the first 4 weeks (precisely why this CamDo effort was a pilot season this year), however we had some small success in the last 2 weeks of deployment that give us hope for the future. The camera recorded a lot of things: thick layers of mysids, countless rockfish and lingcod, several swimming and foraging murres, a handful of harbor seals, and two encounters of the species we were hoping to film – gray whales! While the footage is not the ‘money shot’ we are hoping to film (aka, a headstanding gray whale eating zooplankton right in front of the camera), the fact that we captured gray whales in the first place has showed us that this set-up is a promising investment of time, money and effort that will hopefully deliver next year.

Musings

You may have picked up on the fact that we had slow starts to our field seasons in both Newport and Port Orford. Furthermore, while the number of whale sightings did increase in both locations throughout the field seasons, the number of sightings and whales per day were lower than they have been in previous years. For example, in 2018, we identified 15 different individuals in the month of August in Port Orford (compared to just 5 this year). In 2019, 63 unique whales were seen in Newport (compared to 46 this year). Interestingly, we had a greater diversity of encountered individuals at the start and end of the season in Newport, with a relatively small number of different individuals in July and August. While I cannot provide a definitive reason (or reasons) as to why patterns were observed (we will need to analyze several years of our data to try and understand why), I have some hypotheses I wish to share with you.

As I mentioned in a previous blog, this summer the coastal upwelling along the Oregon coast was delayed (Figure 1). Typically, peak upwelling occurs during the month of June or shortly thereafter, bringing nutrient-rich, deep waters to the surface and, when mixed with sunlight, a lot of productivity. This productivity sets off a chain of reactions — the input of nutrients leads to increased phytoplankton production, which in turn leads to increased zooplankton production, resulting in growth and development of larger organisms that consume zooplankton, such as rockfish and gray whales. If the timing of upwelling is delayed, then so too is this chain of reactions. As you can see from Figure 1, the red lines show that the peak upwelling this year occurred far later in the summer than any year in the last 10 years, with the exception of 2012. Gray whales may have cued into this delay and therefore also delayed their arrival to the PCFG feeding grounds, hence causing us to have low sighting rates at the start of our season. However, this is mostly speculative as we still do not understand the functional mechanisms by which cetaceans, such as gray whales, detect prey across different scales, and to what extent oceanographic conditions like upwelling may play a role in prey availability (Torres 2017). 

Figure 1. 10 year time series of the Coastal Upwelling Transport Index (CUTI). CUTI represents the amount of upwelling (positive numbers) or downwelling (negative numbers). The light-colored lines represent the CUTI at that point in time while the dark, bold line represents the long-term average. The vertical red lines represent the point of peak upwelling in that summer and the horizontal green line shows the peak level of upwelling in 2020 relative to all previous years.

Furthermore, the green line in Figure 1 shows that even after peak upwelling was reached this year, upwelling conditions were lower than all the other peaks in the previous 10 years. We know that weak upwelling is correlated to poor body condition of PCFG gray whales in subsequent years (Soledade Lemos et al. 2020). Upon arriving to the Oregon coast feeding grounds, gray whales may have noticed that it was shaping up to be a poor prey year (we certainly noticed it in Port Orford in the emptiness of our zooplankton net). Faced with this low resource availability, individuals had to make important decisions – risk staying in a currently prey-poor environment or continue the journey onward, searching for better prey conditions elsewhere. This conundrum is known as the marginal value theorem, whereby an individual must decide whether it should abandon the patch it is currently foraging on and move on to search for a new patch without knowing how far away the next patch may be or its value relative to the current patch (Charnov 1976). If we think of the Oregon coast as the ‘current patch’, then we can see how the marginal value theorem translates to the situation gray whales may have found themselves in at the start of the summer. 

Yet, an individual gray whale does not make these decisions in a vacuum. Instead, all gray whales in the same area are faced with the same conundrum. Seminal work by Pianka (1974) showed that when resources, such as food, are abundant, then competition between predators is low because there is enough food to go around. However, when resources dwindle, competition increases and the niches of predators begin to overlap more and more. With Charnov and Pianka’s theories in mind, we can see two groups of gray whales emerge from our 2020 field work observations: those that stayed in the ‘current patch’ (Oregon) and those that decided to seek out a new patch in hopes that it would be a better one. Solé certainly belongs in the first group. We saw her consistently throughout the whole summer. In fact, she was oftentimes so predictable that we would find her foraging on the same reef complex every time we went out to survey. Smudge may also belong in this group, however it is hard to say definitively since we only survey in Port Orford in late July and August. In contrast, I would place whales such as Spray and Heart in the second group since we saw them early in the summer and then not again until mid-to-late September. Where did they go in the interim? Did they go somewhere else in the PCFG range? Or did they venture all the way up to Alaska to the primary Eastern North Pacific (ENP) gray whale feeding grounds? Did their choice to search for food elsewhere pay off?  

As I said earlier, these are all just musings for now, but the GEMM Lab is already hard at work trying to answer these questions. Stay tuned to see what we find!

* Thanks to all the divers who assisted with the pilot CamDo season: Aaron Galloway, Ross Whippo, Svetlana Maslakova, Taylor Eaton, Cori Kane, Austin Williams, Justin Smith

References

Charnov, E.L. 1976. Optimal Foraging, the Marginal Value Theorem. Theoretical Population Biology 9(2):129-136.

Pianka, E.R. 1974. Niche Overlap and Diffuse Competition. PNAS 71(5):2141-2145.

Soledade Lemos, L., Burnett, J.D., Chandler, T.E., Sumich, J.L., and L.G. Torres. 2020. Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11(4):e03094.

Torres, L.G. 2017. A sense of scale: Foraging cetaceans’ use of scale-dependent multimodal sensory systems. Marine Mammal Science 33(4):1170-1193.

Torres, L.G., Nieukirk, S.L., Lemos, L., and T.E. Chandler. 2018. Drone Up! Quantifying Whale Behavior From a New Perspective Improves Observational Capacity. Frontiers in Marine Science: https://doi.org/10.3389/fmars.2018.00319.

Do gray whales count calories?

By Lisa Hildebrand, MSc student, OSU Department of Fisheries & Wildlife, Marine Mammal Institute, Geospatial Ecology of Marine Megafauna Lab

When humans count calories it is typically to regulate and limit calorie intake. What I am wondering about is whether gray whales are aware of caloric differences in the prey that is available to them and whether they make foraging decisions based on those differences. In last week’s post, Dawn discussed what makes a good meal for a hungry blue whale. She discussed that total prey biomass of a patch, as well as how densely aggregated that patch is, are the important factors when a blue whale is picking its next meal. If these factors are important for blue whales, is it same for gray whales? Why even consider the caloric value of their prey?

Gray and blue whales are different in many ways; one way is that blue whales are krill specialists whereas gray whales are more flexible foragers. The Pacific Coast Feeding Group (PCFG) of gray whales in particular are known to pursue a more varied menu. Previous studies along the PCFG range have documented gray whales feeding on mysid shrimp (Darling et al. 1998; Newell 2009), amphipods (Oliver et al. 1984Darling et al. 1998), cumacean shrimp (Jenkinson 2001; Moore et al. 2007; Gosho et al. 2011), and porcelain crab larvae (Dunham and Duffus 2002), to name a few. Based on our observations in the field and from our drone footage, we have observed gray whales feeding on reefs (likely on mysid shrimp), benthically (likely on burrowing amphipods), and at the surface on crab larvae (Fig. 1). Therefore, while both blue and PCFG whales must make decisions about prey patch quality based on biomass and density of the prey, gray whales have an extra decision to make based on prey type since their prey menu items occupy different habitats that require different feeding tactics and amount of energy to acquire them. In light of these reasons, I hypothesize that prey caloric value factors into their decision of prey patch selection. 

Figure 1. Gray whales use several feeding tactics to obtain a variety of coastal Oregon zooplankton prey including jaw snapping (0:12 of video), drooling mud (0:21), and head standing (0:32), to name a few.

This prey selection process is crucial since PCFG gray whales only have about 6 months to consume all the food they need to migrate and reproduce (even less for the Eastern North Pacific (ENP) gray whales since their journey to their Arctic feeding grounds is much longer). You may be asking, well if feeding is so important to gray whales, then why not eat everything they come across? Surely, if they ate every prey item they swam by, then they would be fine. The reason it isn’t quite this simple is because there are energetic costs to travel to, search for, and consume food. If an individual whale simply eats what is closest (a small, poor-quality prey patch) and uses up more energy than it gains, it may be missing out on a much more beneficial and rewarding prey patch that is a little further away (that patch may disperse or another whale may eat it by the time this whale gets there). Scientists have pondered this decision-making process in predators for a long time. These ponderances are best summed up by two central theories: the optimal foraging theory (MacArthur & Pianka 1966) and the marginal value theorem (Charnov 1976). If you are a frequent reader of the blog, you have probably heard these terms once or twice before as a lot of the questions we ask in the GEMM Lab can be traced back to these concepts.

Optimal foraging theory (OFT) states that a predator should pick the most beneficial resource for the lowest cost, thereby maximizing the net energy gained. So, a gray whale should pick a prey patch where it knows that it will gain more energy from consuming the prey in the patch than it will lose energy in the process of searching for and feeding on it. Marginal value theorem elaborates on this OFT concept by adding that the predator also needs to consider the cost of giving up a prey patch to search for a new one, which may or may not end up being more profitable or which may take a very long time to find (and therefore cost more energy). 

The second chapter of my thesis will investigate whether individual gray whales have foraging preferences by relating feeding location to prey quality (community composition) and quantity (relative density). However, in order to do that, I first must know about the quality of the individual prey species, which is why my first chapter explores the caloric content of common coastal zooplankton species in Oregon that may serve as gray whale prey. The lab work and analysis for that chapter are completed and I am in the process of writing it up for publication. Preliminary results (Fig. 2) show variation in caloric content between species (represented by different colors) and reproductive stages (represented by different shapes), with a potential increasing trend throughout the summer. These results suggest that some species and reproductive stages may be less profitable than others based solely on caloric content. 

Figure 2. Mean caloric content (J/mg) of coastal Oregon zooplankton (error bars represent standard deviation) from May-October in 2017-2018. Colors represent species and shapes represent reproductive stage.

Now that we have established that there may be bigger benefits to feeding on some species over others, we have to consider the availability of these zooplankton species to PCFG whales. Availability can be thought of in two ways: 1) is the prey species present and at high enough densities to make searching and foraging profitable, and 2) is the prey species in a habitat or depth that is accessible to the whale at a reasonable energetic cost? Some prey species, such as crab larvae, are not available at all times of the summer. Their reproductive cycles are pulsed (Roegner et al. 2007) and therefore these prey species are less available than species, such as mysid shrimp, that have more continuous reproduction (Mauchline 1980). Mysid shrimp appear to seek refuge on reefs in rock crevices and among kelp, whereas amphipods often burrow in soft sediment. Both of these habitat types present different challenges and energetic costs to a foraging gray whale; it may take more time and energy to dislodge mysids from a reef, but the payout will be bigger in terms of caloric gain than if the whale decides to sift through soft sediment on the seafloor to feed on amphipods. This benthic feeding tactic may potentially be a less costly foraging tactic for PCFG whales, but the reward is a less profitable prey item.  

My first chapter will extend our findings on the caloric content of Oregon coastal zooplankton to facilitate a comparison to the caloric values of the main ampeliscid amphipod prey of ENP gray whales feeding in the Arctic. Through this comparison I hope to assess the trade-offs of being a PCFG whale rather than an ENP whale that completes the full migration cycle to the primary summer feeding grounds in the Arctic. 

References

Charnov, E. L. 1976. Optimal foraging: the marginal value theorem. Theoretical Population Biology 9:129-136.

Darling, J. D., Keogh, K. E. and T. E. Steeves. 1998. Gray whale (Eschrichtius robustus) habitat utilization and prey species off Vancouver Island, B.C. Marine Mammal Science 14(4):692-720.

Dunham, J. S. and D. A. Duffus. 2002. Diet of gray whales (Eschrichtius robustus) in Clayoquot Sound, British Columbia, Canada. Marine Mammal Science 18(2):419-437.

Gosho, M., Gearin, P. J., Jenkinson, R. S., Laake, J. L., Mazzuca, L., Kubiak, D., Calambokidis, J. C., Megill, W. M., Gisborne, B., Goley, D., Tombach, C., Darling, J. D. and V. Deecke. 2011. SC/M11/AWMP2 submitted to International Whaling Commission Scientific Committee.

Jenkinson, R. S. 2001. Gray whale (Eschrichtius robustus) prey availability and feeding ecology in Northern California, 1999-2000. Master’s thesis, Humboldt State University.

MacArthur, R. H., and E. R. Pianka. 1966. On optimal use of a patchy environment. American Naturalist 100:603-609.

Mauchline, J. 1980. The larvae and reproduction in Blaxter, J. H. S., Russell, F. S., and M. Yonge, eds. Advances in Marine Biology vol. 18. Academic Press, London.

Moore, S. E., Wynne, K. M., Kinney, J. C., and C. M. Grebmeier. 2007. Gray whale occurrence and forage southeast of Kodiak Island, Alaska. Marine Mammal Science 23(2)419-428.

Newell, C. L. 2009. Ecological interrelationships between summer resident gray whales (Eschrichtius robustus) and their prey, mysid shrimp (Holmesimysis sculpta and Neomysis rayii) along the central Oregon coast. Master’s thesis, Oregon State University.

Oliver, J. S., Slattery, P. N., Silberstein, M. A., and E. F. O’Connor. 1984. Gray whale feeding on dense ampeliscid amphipod communities near Bamfield, British Columbia. Canadian Journal of Zoology 62:41-49.

Roegner, G. C., Armstrong, D. A., and A. L. Shanks. 2007. Wind and tidal influences on larval crab recruitment to an Oregon estuary. Marine Ecology Progress Series 351:177-188.