Are there picky eaters in the PCFG?

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

As anyone who has ever been, or raised, a picky eater knows, humans have a wide range of food preferences. The diversity of available cuisines is a testament to the fact that we have individual food preferences. While taste is certainly a primary influence, nutritional benefits and accessibility are other major factors that affect our eating choices. But we are not the only species to have food preferences. In cetacean research, it is common to study the prey types consumed by a population as a whole. Narrowing these prey preferences down to the individual level is rare. While the individual component is challenging to study and to incorporate into population models, it is important to consider what the effects of individual foraging specialization might be.

To understand the role and drivers of individual specialization in population ecology, it is important to first understand the concepts of niche variation and partitioning. An animal’s ecological niche describes its role in the ecosystem it inhabits (Hutchinson, 1957). A niche is multidimensional, with dimensions for different environmental conditions and resources that a species requires. One focus of my research pertains to the dimensions of the niche related to foraging. As discussed in a previous blog, niche partitioning occurs when ecological space is shared between competitors through access to resources varies across different dimensions such as prey type, foraging location, and time of day when foraging takes place. Niche partitioning is usually discussed on the scale of different species coexisting in an ecosystem. Pianka’s theory stating that niche partitioning will increase as prey availability decreases uses competing lizard species as the example (Pianka, 1974). Typically, niche partitioning theory considers inter-specific competition (competition between species), but niche partitioning can take place within a species in response to intra-specific competition (competition between individuals of the same species) through individual niche variation.

A species that consumes a multitude of prey types is considered a generalist while one with a specific prey type is considered a specialist. Gray whales are considered generalists (Nerini, 1984). However, we do not know if each individual gray whale is a generalist or if the generalist population is actually composed of individual specialists with different preferences. One way to test for the presence of individual specialization is to compare the niche width of the population to the niche width of each individual (Figure 1, Bolnick et al., 2003).  For example, if a population eats five different types of prey and each individual consumed those prey types, those individuals would be generalists. However, if each individual only consumed one of the prey types, then those individuals would be specialists within a generalist population.

Figure 1. Figure from Bolnick et al. 2003. The thick curve represents the total niche of the population and the thin curves represent individual niches. Note that in both panels the population has the same total niche. In panel A, the individual curves overlap and are all pretty wide. These curves represent individual generalists that make up a generalist population. In panel B, the thin curves are narrower and do not overlap as much as those in panel A. These curves represent individual specialists that make up a generalist population.

If individual specialization is present in a population the natural follow-up question is why? To answer this, we look for common characteristics between the individuals that are similarly specialized. What do all the individuals that feed on the same prey type have in common? Common characterizations that may be found include age, sex, or distinct morphology (such as different beak or body shapes) (Bolnick et al., 2003).

Woo et al. (2008) studied individual specialization in Brünnich’s guillemot, a generalist sea bird species, using diet and tagging data. They found individual specialization in both diet (prey type) and behavior (dive depth, shape, and flight time). Specialization occurred across multiple timescales but was higher over short-time scales. The authors found that it was more common for an individual to specialize in a prey-type/foraging tactic for a few days than for that specialization to continue across years, although a few individuals were specialists for the full 15-year period of the study. Based on reproductive success of the studies birds, the authors concluded that the generalist and specialist strategies were largely equivalent in terms of fitness and survival. The authors searched for common characteristics in the individuals with similar specialization and they found that the differences between sexes or age classes were so small that neither grouping explained the observed individual specialization. This is an interesting result because it suggests that there is some missing attribute, that of the authors did not examine, that might explain why individual specialists were present in the population.

Hoelzel et al. (1989) studied minke whale foraging specialization by observing the foraging behaviors of 23 minke whales over five years from a small boat. They identified two foraging tactics: lunge feeding and bird-associated feeding. Lunge feeding involved lunging up through the water with an open mouth to engulf a group of fish, while bird-associated feeding took advantage of a group of fish being preyed on by sea birds to attack the fish from below while they were already being attacked from above. They found that nine individuals used lunge feeding, and of those nine, six whales used this tactic exclusively. Five of those six whales were observed in at least two years. Seventeen whales were observed using bird-associated feeding, 14 exclusively. Of those 14, eight were observed in at least two years. Interestingly, like Woo et al. (2008), this study did not find any associations between foraging tactic use and sex, age, or size of whale. Through a comparison of dive durations and feeding rates, they hypothesized that lunge feeding was more energetically costly but resulted in more food, while bird-associated feeding was energetically cheaper but had a lower capture rate. This result means that these two strategies might have the similar energetic payoffs.

Both of these studies are examples of questions that I am excited to ask using our data on the PCFG gray whales feeding off the Oregon coast (especially after doing the research for this blog). We have excellent individual-specific data to address questions of specialization because the field teams for  this project always carefully link observed behaviors with individual whale ID.  Using these data, I am curious to find out if the whales in our study group are individual specialists or generalists (or some combination of the two). I am also interested in relating specific tactics to their energetic costs and benefits in order to assess the payoffs of each foraging tactic. I then hope to combine the results of both analyses to assess the payoffs of each individual whale’s strategy.

Figure 2. Example images of two foraging tactics, side swimming (left) and headstanding (right). Images captured under NOAA/NMFS permit #21678.

Studying individual specialization is important for conservation. Consider the earlier example of a generalist population that consumes five prey items but is composed of individual specialists. If the presence of individual specialization is not accounted for in management plans, then regulations may protect certain prey types or foraging tactics/areas of the whales and not others. Such a management plan could be a dangerous outcome for the whale population because only parts of the population would be protected, while other specialists are at risk, thus potentially losing genetic diversity, cultural behaviors, and ecological resilience in the population as a whole. A plan designed to maximize protection for all the specialists would be better for the population because populations with increased ecological resilience are more likely to persist through periods of rapid environmental change. Furthermore, understanding individual specialization could help us better predict how a population might be affected by environmental change. Environmental change does not affect all prey species in the same way. An individual specialization study could help identify which whales might be most affected by predicted environmental changes. Therefore, in addition to being a fascinating and exciting research question, it is important to test for individual specialization in order to improve management and our overall understanding of the PCFG gray whale population.

References

Bolnick, D. I., Svanbäck, R., Fordyce, J. A., Yang, L. H., Davis, J. M., Hulsey, C. D., & Forister, M. L. (2003). The ecology of individuals: Incidence and implications of individual specialization. American Naturalist, 161(1), 1–28. https://doi.org/10.1086/343878

Hoelzel, A. R., Dorsey, E. M., & Stern, S. J. (1989). The foraging specializations of individual minke whales. Animal Behaviour, 38(5), 786–794. https://doi.org/10.1016/S0003-3472(89)80111-3

Hutchinson, G. E. (1957). Concluding Remarks. Cold Spring Harbor Symposia on Quantitative Biology, 22(0), 415–427. https://doi.org/10.1101/sqb.1957.022.01.039

Nerini, M. (1984). A Review of Gray Whale Feeding Ecology. In The Gray Whale: Eschrichtius Robustus (pp. 423–450). Elsevier Inc. https://doi.org/10.1016/B978-0-08-092372-7.50024-8

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

Woo, K. J., Elliott, K. H., Davidson, M., Gaston, A. J., & Davoren, G. K. (2008). Individual specialization in diet by a generalist marine predator reflects specialization in foraging behaviour. Journal of Animal Ecology, 77(6), 1082–1091. https://doi.org/10.1111/j.1365-2656.2008.01429.x

Why Feeling Stupid is Great: How stupidity fuels scientific progress and discovery

By Alexa Kownacki, Ph.D. Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

It all started with a paper. On Halloween, I sat at my desk, searching for papers that could answer my questions about bottlenose dolphin metabolism and realized I had forgotten to check my email earlier. In my inbox, there was a new message with an attachment from Dr. Leigh Torres to the GEMM Lab members, saying this was a “must-read” article. The suggested paper was Martin A. Schwartz’s 2008 essay, “The importance of stupidity in scientific research”, published in the Journal of Cell Science, highlighted universal themes across science. In a single, powerful page, Schwartz captured my feelings—and those of many scientists: the feeling of being stupid.

For the next few minutes, I stood at the printer and absorbed the article, while commenting out loud, “YES!”, “So true!”, and “This person can see into my soul”. Meanwhile, colleagues entered my office to see me, dressed in my Halloween costume—as “Amazon’s Alexa”, talking aloud to myself. Coincidently, I was feeling pretty stupid at that moment after just returning from a weekly meeting, where everyone asked me questions that I clearly did not have the answers to (all because of my costume). This paper seemed too relevant; the timing was uncanny. In the past few weeks, I have been writing my PhD research proposal —a requirement for our department— and my goodness, have I felt stupid. The proposal outlines my dissertation objectives, puts my work into context, and provides background research on common bottlenose dolphin health. There is so much to know that I don’t know!

Alexa dressed as “Amazon Alexa” on Halloween at her office in San Diego, CA.

When I read Schwartz’s 2008 paper, there were a few takeaway messages that stood out:

  1. People take different paths. One path is not necessarily right nor wrong. Simply, different. I compared that to how I split my time between OSU and San Diego, CA. Spending half of the year away from my lab and my department is incredibly challenging; I constantly feel behind and I miss the support that physically being with other students provides. However, I recognize the opportunities I have in San Diego where I work directly with collaborators who teach and challenge me in new ways that bring new skills and perspective.

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    (Image source: St. Albert’s Place)
  2. Feeling stupid is not bad. It can be a good feeling—or at least we should treat it as being a positive thing. It shows we have more to learn. It means that we have not reached our maximum potential for learning (who ever does?). While writing my proposal I realized just how little I know about ecotoxicology, chemistry, and statistics. I re-read papers that are critical to understanding my own research, like “Nontargeted biomonitoring of halogenated organic compounds in two ecotypes of bottlenose dolphins (Tursiops truncatus) from the Southern California bight” (2014) by Shaul et al. and “Bottlenose dolphins as indicators of persistent organic pollutants in the western north Atlantic ocean and northern gulf of Mexico” (2011) by Kucklick et al. These articles took me down what I thought were wormholes that ended up being important rivers of information. Because I recognized my knowledge gap, I can now articulate the purpose and methods of analysis for specific compounds that I will conduct using blubber samples of common bottlenose dolphins

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    Image source: memegenerator.net
  3. Drawing upon experts—albeit intimidating—is beneficial for scientific consulting as well as for our mental health; no one person knows everything. That statement can bring us together because when people work together, everyone benefits. I am also reminded that we are our own harshest critics; sometimes our colleagues are the best champions of our own successes. It is also why historical articles are foundational. In the hunt for the newest technology and the latest and greatest in research, it is important to acknowledge the basis for discoveries. My data begins in 1981, when the first of many researchers began surveying the California coastline for common bottlenose dolphins. Geographic information systems (GIS) were different back then. The data requires conversions and investigative work. I had to learn how the data were collected and how to interpret that information. Therefore, it should be no surprise that I cite literature from the 1970s, such as “Results of attempts to tag Atlantic Bottlenose dolphins, (Tursiops truncatus)” by Irvine and Wells. Although published in 1972, the questions the authors tried to answer are very similar to what I am looking at now: how are site fidelity and home ranges impacted by natural and anthropogenic processes. While Irvine and Wells used large bolt tags to identify individuals, my project utilizes much less invasive techniques (photo-identification and blubber biopsies) to track animals, their health, and their exposures to contaminants.

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    (Image source: imgflip.com)
  4. Struggling is part of the solution. Science is about discovery and without the feeling of stupidity, discovery would not be possible. Feeling stupid is the first step in the discovery process: the spark that fuels wanting to explore the unknown. Feeling stupid can lead to the feeling of accomplishment when we find answers to those very questions that made us feel stupid. Part of being a student and a scientist is identifying those weaknesses and not letting them stop me. Pausing, reflecting, course correcting, and researching are all productive in the end, but stopping is not. Coursework is the easy part of a PhD. The hard part is constantly diving deeper into the great unknown that is research. The great unknown is simultaneously alluring and frightening. Still, it must be faced head on. Schwartz describes “productive stupidity [as] being ignorant by choice.” I picture this as essentially blindly walking into the future with confidence. Although a bit of an oxymoron, it resonates the importance of perseverance and conviction in the midst of uncertainty.

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    (Image source: Redbubble)

Now I think back to my childhood when stupid was one of the forbidden “s-words” and I question whether society had it all wrong. Maybe we should teach children to acknowledge ignorance and pursue the unknown. Stupid is a feeling, not a character flaw. Stupidity is important in science and in life. Fascination and emotional desires to discover new things are healthy. Next time you feel stupid, try running with it, because more often than not, you will learn something.

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Alexa teaching about marine mammals to students ages 2-6 and learning from educators about new ways to engage young students. San Diego, CA in 2016. (Photo source: Lori Lowder)