## Species distribution modeling: Part statistics, part philosophy, and there is no “right answer”

By Dawn Barlow, PhD student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

Just like that, I have wrapped up year 1 of my PhD in Wildlife Science. For my PhD, I am investigating the ecology and distribution of blue whales in New Zealand across multiple spatial and temporal scales. In a region where blue whales overlap with industrial activity, there is considerable interest from managers to be able to reliably forecast when and where blue whales are most likely to be in the area. In a series of five chapters and utilizing multiple different data sources (dedicated boat surveys, oceanographic data, acoustic recordings, remotely sensed environmental data, opportunistic blue whale sightings information), I will attempt to describe, quantify, and predict where blue whales are found in relation to their environment. Each chapter will evaluate the distribution of blue whales relative to the environment at different scales in space (ranging from 4 km to 25 km resolution) and time (ranging from daily to seasonal resolution). One overarching method I am using throughout my PhD is species distribution modeling. Having just completed my research review with my doctoral committee last week, I’ll share this aspect of my research proposal that I’ve particularly enjoyed reading, writing, and thinking about.

Species distribution models (SDMs), which are sometimes referred to as habitat models or ecological niche models, are mathematical algorithms that combine observations of a species with environmental conditions at their observed locations, to gain ecological insight and predict spatial distributions of the species (Elith and Leathwick, 2009; Redfern et al., 2006). Any model is just one description of what is occurring in the natural world. Just as there are many ways to describe something with words and many languages to do so, there are many options for modeling frameworks and approaches, with stark and nuanced differences. My labmate and friend Solene Derville has equated the number of choices one has for SDMs to the cracker section in an American grocery store. When navigating all of these choices and considerations, it is important to remember that no model will ever be completely correct—it is our best attempt at describing a complex natural system—and as an analyst we need to do the best that we can with the data available to address the ecological questions at hand. As it turns out, the dividing line between quantitative analysis and philosophy is thin at times. What may seem at first like a purely objective, statistical endeavor requires careful consideration and fundamental decision-making on the part of the analyst.

Ecosystems are multifaceted, complex, and hierarchical. They are comprised of multiple physical and biological components, which operate at multiple scales across space and time. As Dr. Simon Levin stated in at 1989 MacArthur Award lecture on the topic of scale in ecology:

“A good model does not attempt to reproduce every detail of the biological system; the system itself suffices for that purpose as the most detailed model of itself. Rather, the objective of a model should be to ask how much detail can be ignored without producing results that contradict specific sets of observations, on particular scales of interest” (Levin, 1992).

The question of scale is central to ecology. As many biology students learn in their first introductory classes, parsimony is “The principle that the most acceptable explanation of an occurrence, phenomenon, or event is the simplest, involving the fewest entities, assumptions, or changes” (Oxford Dictionary). In other words, the best explanation is the simplest one. One challenge in ecological modeling, including SDMs, is to select spatial and temporal scales as coarse as possible for the most parsimonious—the most straightforward—model, while still being fine enough to capture relevant patterns. Another critical consideration is the scale of the question you are interested in answering. The scale of the analysis must match the scale at which you want to make inferences about the ecology of a species.

Similarly, the issue of complexity is central to distribution modeling. Overly simple models may not be able to adequately describe the relationship between species occurrence and the environment. In contrast, highly complex models may have very high explanatory power, but risk ascribing an ecological pattern to noise in the data (Merow et al., 2014), in other words, finding patterns that aren’t real. Furthermore, highly complex models tend to have poorer predictive capacity than simpler models (Merow et al., 2014). There is a trade-off between descriptive and predictive power in SDMs (Derville et al., 2018). Therefore, a key component in the SDM process is establishing the end goal of the model with respect to the region of interest, scale, explanatory power, predictive capacity, and in many cases management need.

Finally, any model is ultimately limited by the data available and the scale at which it was collected (Elith and Leathwick, 2009; Guillera-Arroita et al., 2015; Redfern et al., 2006). Prior knowledge of what environmental features are important to the species of interest is often limited at the time of the data collection effort, and data collection is constrained by when it is logistically feasible to sample. For example, we collect detailed oceanographic data during the summer months when it is practical to get out on the water, satellite imagery of sea surface temperature might be unavailable during times of cloud cover, and people are more likely to report blue whale sightings in areas where there is more human activity. Therefore, useful SDMs that address both ecological and management needs typically balance the scale of analysis and model complexity with the limitations of the data.

Managers and politicians within the New Zealand government are interested in a tool to predict when and where blue whales are most likely to be, based on sound ecological analysis. This is one of the end-goals of my PhD, but in the meantime, I am grappling with the appropriate scales of analysis, and attempting to balance questions of model complexity, explanatory power, and predictive capacity. There is no single, correct answer, and so my process is in part quantitative analysis, part philosophy, and all with the goal of increased ecological understanding and conservation of a species.

References:

Derville, S., Torres, L. G., Iovan, C., and Garrigue, C. (2018). Finding the right fit: Comparative cetacean distribution models using multiple data sources and statistical approaches. Divers. Distrib. 24, 1657–1673. doi:10.1111/ddi.12782.

Elith, J., and Leathwick, J. R. (2009). Species Distribution Models: Ecological Explanation and Prediction Across Space and Time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697. doi:10.1146/annurev.ecolsys.110308.120159.

Guillera-Arroita, G., Lahoz-Monfort, J. J., Elith, J., Gordon, A., Kujala, H., Lentini, P. E., et al. (2015). Is my species distribution model fit for purpose? Matching data and models to applications. Glob. Ecol. Biogeogr. 24, 276–292. doi:10.1111/geb.12268.

Levin, S. A. (1992). The problem of pattern and scale. Ecology 73, 1943–1967.

Merow, C., Smith, M. J., Edwards, T. C., Guisan, A., Mcmahon, S. M., Normand, S., et al. (2014). What do we gain from simplicity versus complexity in species distribution models? Ecography (Cop.). 37, 1267–1281. doi:10.1111/ecog.00845.

Redfern, J. V., Ferguson, M. C., Becker, E. A., Hyrenbach, K. D., Good, C., Barlow, J., et al. (2006). Techniques for cetacean-habitat modeling. Mar. Ecol. Prog. Ser. 310, 271–295. doi:10.3354/meps310271.

## Zooming in: A closer look at bottlenose dolphin distribution patterns off of San Diego, CA

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

Data analysis is often about parsing down data into manageable subsets. My project, which spans 34 years and six study sites along the California coast, requires significant data wrangling before full analysis. As part of a data analysis trial, I first refined my dataset to only the San Diego survey location. I chose this dataset for its standardization and large sample size; the bulk of my sightings, over 4,000 of the 6,136, are from the San Diego survey site where the transect methods were highly standardized. In the next step, I selected explanatory variable datasets that covered the sighting data at similar spatial and temporal resolutions. This small endeavor in analyzing my data was the first big leap into understanding what questions are feasible in terms of variable selection and analysis methods. I developed four major hypotheses for this San Diego site.

#### Hypotheses:

H1: I predict that bottlenose dolphin sightings along the San Diego transect throughout the years 1981-2015 exhibit clustered distribution patterns as a result of the patchy distributions of both the species’ preferred habitats, as well as the social nature of bottlenose dolphins.

H2: I predict there would be higher densities of bottlenose dolphin at higher latitudes spanning 1981-2015 due to prey distributions shifting northward and less human activities in the northerly sections of the transect.

H3: I predict that during warm (positive) El Niño Southern Oscillation (ENSO) months, the dolphin sightings in San Diego would be distributed more northerly, predominantly with prey aggregations historically shifting northward into cooler waters, due to (secondarily) increasing sea surface temperatures.

H4: I predict that along the San Diego coastline, bottlenose dolphin sightings are clustered within two kilometers of the six major lagoons, with no specific preference for any lagoon, because the murky, nutrient-rich waters in the estuarine environments are ideal for prey protection and known for their higher densities of schooling fishes.

#### Data Description:

The common bottlenose dolphin (Tursiops truncatus) sighting data spans 1981-2015 with a few gap years. Sightings cover all months, but not in all years sampled. The same transect in San Diego was surveyed in a small, rigid-hulled inflatable boat with approximately a two-kilometer observation area (one kilometer surveyed 90 degrees to starboard and port of the bow).

I wanted to see if there were changes in dolphin distribution by latitude and, if so, whether those changes had a relationship to ENSO cycles and/or distances to lagoons. For ENSO data, I used the NOAA database that provides positive, neutral, and negative indices (1, 0, and -1, respectively) by each month of each year. I matched these ENSO data to my month-date information of dolphin sighting data. Distance from each lagoon was calculated for each sighting.

#### Results:

H1: True, dolphins are clustered and do not have a uniform distribution across this area. Spatial analysis indicated a less than a 1% likelihood that this clustered pattern could be the result of random chance (Fig. 1, z-score = -127.16, p-value < 0.0001). It is well-known that schooling fishes have a patchy distribution, which could influence the clustered distribution of their dolphin predators. In addition, bottlenose dolphins are highly social and although pods change in composition of individuals, the dolphins do usually transit, feed, and socialize in small groups.

H2: False, dolphins do not occur at higher densities in the higher latitudes of the San Diego study site. The sightings are more clumped towards the lower latitudes overall (p < 2e-16), possibly due to habitat preference. The sightings are closer to beaches with higher human densities and human-related activities near Mission Bay, CA. It should be noted, that just north of the San Diego transect is the Camp Pendleton Marine Base, which conducts frequent military exercises and could deter animals.

H3: False, during warm (positive) El Niño Southern Oscillation (ENSO) months, the dolphin sightings in San Diego were more southerly. In colder (negative) ENSO months, the dolphins were more northerly. The differences between sighting latitude and ENSO index was significant (p<0.005). Post-hoc analysis indicates that the north-south distribution of dolphin sightings was different during each ENSO state.

H4: True, dolphins are clustered around particular lagoons. Figure 5 illustrates how dolphin sightings nearest to Lagoon 6 (the San Dieguito Lagoon) are always within 0.03 decimal degrees. Because of how these data are formatted, decimal degrees is the easiest way to measure change in distance (in this case, the difference in latitude). In comparison, dolphins at Lagoon 5 (Los Penasquitos Lagoon) are distributed across distances, with the most sightings further from the lagoon.

I found a significant difference between distance to nearest lagoon in different ENSO index categories (p < 2.55e-9): there is a significant difference in distance to nearest lagoon between neutral and negative values and positive and neutral years. Therefore, I hypothesize that in neutral ENSO months compared to positive and negative ENSO months, prey distributions are changing. This is one possible hypothesis for the significant difference in lagoon preference based on the monthly ENSO index. Using a violin plot (Fig. 6), it appears that Lagoon 5, Los Penasquitos Lagoon, has the widest variation of sighting distances in all ENSO index conditions. In neutral years, Lagoon 0, the Buena Vista Lagoon has multiple sightings, when in positive and negative years it had either no sightings or a single sighting. The Buena Vista Lagoon is the most northerly lagoon, which may indicate that in neutral ENSO months, dolphin pods are more northerly in their distribution.

#### Takeaways to science and management:

Bottlenose dolphins have a clustered distribution which seems to be related to ENSO monthly indices, and likely, their social structures. From these data, neutral ENSO months appear to have something different happening compared to positive and negative months, that is impacting the sighting distributions of bottlenose dolphins off the San Diego coastline. More research needs to be conducted to determine what is different about neutral months and how this may impact this dolphin population. On a finer scale, the six lagoons in San Diego appear to have a spatial relationship with dolphin sightings. These lagoons may provide critical habitat for bottlenose dolphins and/or for their preferred prey either by protecting the animals or by providing nutrients. Different lagoons may have different spans of impact, that is, some lagoons may have wider outflows that create larger nutrient plumes.

Other than the Marine Mammal Protection Act and small protected zones, there are no safeguards in place for these dolphins, whose population hovers around 500 individuals. Therefore, specific coastal areas surrounding lagoons that are more vulnerable to habitat loss, habitat degradation, and/or are more frequented by dolphins, may want greater protection added at a local, state, or federal level. For example, the Batiquitos and San Dieguito Lagoons already contain some Marine Conservation Areas with No-Take Zones within their reach. The city of San Diego and the state of California need better ways to assess the coastlines in their jurisdictions and how protecting the marine, estuarine, and terrestrial environments near and encompassing the coastlines impacts the greater ecosystem.

This dive into my data was an excellent lesson in spatial scaling with regards to parsing down my data to a single study site and in matching my existing data sets to other data that could help answer my hypotheses. Originally, I underestimated the robustness of my data. At first, I hesitated when considering reducing the dolphin sighting data to only include San Diego because I was concerned that I would not be able to do the statistical analyses. However, these concerns were unfounded. My results are strongly significant and provide great insight into my questions about my data. Now, I can further apply these preliminary results and explore both finer and broader scale resolutions, such as using the more precise ENSO index values and finding ways to compare offshore bottlenose dolphin sighting distributions.

## Midway Atoll: the next two weeks at the largest albatross colony in the world (two years later)

By Rachael Orben, Assistant Professor (Senior Research), Seabird Oceanography Lab

This February I had the opportunity to spend two weeks at Midway Atoll National Wildlife Refuge in the Papahānaumokuākea Marine National Monument. I was there to GPS track black-footed and Laysan albatross during their short chick-brooding foraging trips. Two weeks is just enough time since the albatross are taking short trips (3-5 days) to feed their rapidly growing chicks.

My first visit to Midway (2016 blog post) occurred right as the black-footed albatross chicks were hatching (quickly followed by the Laysan albatross chicks). This time, we arrived almost exactly when I had left off. The oldest chicks were just about two weeks old. This shift in phenology meant that, though subtle, each day offered new insights for me as I watched chicks transform into large aware and semi-mobile birds. By the time we left, unattended chicks were rapidly multiplying as the adults shifted to the chick-rearing stage. During chick rearing, both parents leave the chick unattended and take longer foraging trips.

Our research goal was to collect tracking data from both species that can be used to address a couple of research questions. First of all, winds can aid, or hinder albatross foraging and flight efficiency (particularly during the short brooding trips). In the North Pacific, the strength and direction of the winds are influenced by the ENSO (El Niño Southern Oscillation) cycles. The day after we left Midway, NOAA issued an El Niño advisory indicating weak El Nino conditions. We know from previous work at Tern Island (farther east and farther south at 23.87 N, -166.28 W) that El Niño improves foraging for Laysan albatrosses during chick brooding, while during La Niña reproductive success is lower (Thorne et al., 2016). However, since Midway is farther north, and farther west the scenario might be different there. Multiple years of GPS tracking data are needed to address this question and we hope to return to collect more data next year (especially if  La Niña follows the El Niño as is often the case).

We will also overlap the tracking data with fishing boat locations from the Global Fishing Watch database to assess the potential for birds from Midway to interact with high seas fisheries during this time of year (project description, associated blog post). Finally, many of the tags we deployed incorporated a barometric pressure sensor and the data can be used to estimate flight heights relative to environmental conditions such as wind strength. This type of data is key to assessing the impact of offshore wind energy (Kelsey et al., 2018).

#### How to track an albatross

To track an albatross we use small GPS tags that we tape to the back feathers. After the bird returns from a foraging trip, we remove the tape from the feathers and take the datalogger off. Then we recharge the battery and download the data!

This research is a collaboration between Lesley Thorne (Stony Brook University), Scott Shaffer (San Jose State University), myself (Oregon State University), and Melinda Conners (Washington State University). The field effort was generously supported by the Laurie Landeau Foundation via the Minghua Zhang Early Career Faculty Innovation Fund at Stoney Brook University to Lesley Thorne.

`My previous visit to Midway occurred just after house mice were discovered attacking incubating adult albatrosses. Since then, a lot of thought and effort had gone into developing a plan to eradicate mice from Midway. You can find out more via Island Conservation’s Midway blogs and the USFWS.`
###### References

Kelsey, E. C., Felis, J. J., Czapanskiy, M., Pereksta, D. M., & Adams, J. (2018). Collision and displacement vulnerability to offshore wind energy infrastructure among marine birds of the Pacific Outer Continental Shelf. Journal of Environmental Management, 227, 229–247. http://doi.org/10.1016/j.jenvman.2018.08.051

Thorne, L. H., Conners, M. G., Hazen, E. L., Bograd, S. J., Antolos, M., Costa, D. P., & Shaffer, S. A. (2016). Effects of El Niño-driven changes in wind patterns on North Pacific albatrosses. Journal of the Royal Society Interface, 13(119), 20160196. http://doi.org/10.1098/rsif.2016.0196

## More than just whales: The importance of studying an ecosystem

By Dawn Barlow, PhD student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

I have the privilege of studying the largest animals on the planet: blue whales (Balaenoptera musculus). However, in order to understand the ecology, distribution, and habitat use patterns of these ocean giants, I have dedicated the past several months to studying something much smaller: krill (Nyctiphanes australis). New Zealand’s South Taranaki Bight region (“STB”, Figure 1) is an important foraging ground for a unique population of blue whales [1,2]. A wind-driven upwelling system off of Kahurangi Point (the “X” in Figure 1) generates productivity in the region [3], leading to an abundance of krill [4], the desired blue whale prey [5].

Our blue whale research team collected a multitude of datastreams in three different years, including hydroacoustic data to map krill distribution throughout our study region. The summers of 2014 and 2017 were characterized by what could be considered “typical” conditions: A plume of cold, upwelled water curving its way around Cape Farewell (marked with the star in Figure 1) and entering the South Taranaki Bight, spurring a cascade of productivity in the region. The 2016 season, however, was different. The surface water temperatures were hot, and the whales were not where we expected to find them.

What happened to the blue whales’ food source under these different conditions in 2016? Before I share some preliminary findings from my recent analyses, it is important to note that there are many possible ways to measure krill availability. For example, the number of krill aggregations, as well as how deep, thick, and dense those aggregations are in an area will all factor into how “desirable” krill patches are to a blue whale. While there may not be “more” or “less” krill from one year to the next, it may be more or less accessible to a blue whale due to energetic costs of capturing it. Here is a taste of what I’ve found so far:

In 2016, when surface waters were warm, the krill aggregations were significantly deeper than in the “typical” years (ANOVA, F=7.94, p <0.001):

The number of aggregations was not significantly different between years, but as you can see in the plot below (Figure 4) the krill were distributed differently in space:

While the bulk of the krill aggregations were located north of Cape Farewell under typical conditions (2014 and 2017), in the warm year (2016) the krill were not in this area. Rather, the area with the most aggregations was offshore, in the western portion of our study region. Now, take a look at the same figure, overlaid with our blue whale sighting locations:

Where did we find the whales? In each year, most whale encounters were in the locations where the most krill aggregations were found! Not only that, but in 2016 the whales responded to the difference in krill distribution by shifting their distribution patterns so that they were virtually absent north of Cape Farewell, where most sightings were made in the typical years.

The above figures demonstrate the importance of studying an ecosystem. We could puzzle and speculate over why the blue whales were further west in the warm year, but the story that is emerging in the krill data may be a key link in our understanding of how the ecosystem responds to warm conditions. While the focus of my dissertation research is blue whales, they do not live in isolation. It is through understanding the ecosystem-scale story that we can better understand blue whale ecology in the STB. As I continue modeling the relationships between oceanography, krill, and blue whales in warm and typical years, we are beginning to scratch the surface of how blue whales may be responding to their environment.

1. Torres LG. 2013 Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zeal. J. Mar. Freshw. Res. 47, 235–248. (doi:10.1080/00288330.2013.773919)
2. Barlow DR et al. 2018 Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger. Species Res. 36, 27–40. (doi:https://doi.org/10.3354/esr00891)
3. Shirtcliffe TGL, Moore MI, Cole AG, Viner AB, Baldwin R, Chapman B. 1990 Dynamics of the Cape Farewell upwelling plume, New Zealand. New Zeal. J. Mar. Freshw. Res. 24, 555–568. (doi:10.1080/00288330.1990.9516446)
4. Bradford-Grieve JM, Murdoch RC, Chapman BE. 1993 Composition of macrozooplankton assemblages associated with the formation and decay of pulses within an upwelling plume in greater cook strait, New Zealand. New Zeal. J. Mar. Freshw. Res. 27, 1–22. (doi:10.1080/00288330.1993.9516541)
5. Gill P. 2002 A blue whale (Balaenoptera musculus) feeding ground in a southern Australian coastal upwelling zone. J. Cetacean Res. Manag. 4, 179–184.

## Hundreds and hundreds and hundreds of models: An ecologist’s love for programming

By Dawn Barlow, PhD student, Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

When people hear that I study blue whales, they often ask me questions about what it’s like to be close to the largest animal on the planet, where we do fieldwork, and what data we are interested in collecting. While I love time at sea, my view on a daily basis is rarely like this:

More often than not, it looks something like this:

In my application letter to Dr. Leigh Torres, I wrote something along the lines of “while I relish remote fieldwork, I also find great satisfaction in the analysis process.” This statement is increasingly true for me as I grow more proficient in statistical modeling and computer programming. When excitedly telling my family about how I am trying to model relationships between oceanography, krill, whales, and satellite imagery, I was asked what I meant by “model”. Put simply, a model is a formula or equation that we can use to describe a pattern. I have been told, “all models are wrong, but some models work.” What does this mean? While we may never know exactly every pattern of whale feeding behavior, we can use the data we have to describe some of the important relationships. If our model performance is very good, then we have likely described most of what drives the patterns we see. If model performance is poor, then there is more to the pattern that we have not yet captured in either our data collection or in our analytical methods. Another common saying about models is, “A model is only ever as good as the data you put into it.” While we worked hard during field seasons to collect a myriad of data about what could be influencing blue whale distribution patterns, we inevitably could not capture everything, nor do we know everything that should be measured.

So, how do you go about finding the ‘best’ model? This question is what I’ve been grappling with over the last several weeks. My goal is to describe the patterns in the krill that drive patterns in whale distribution, the patterns in oceanography that drive patterns in the krill, and the patterns in the oceanography that drive patterns in whale distribution. The thing is, we have many metrics to describe oceanographic patterns (surface temperature, mixed layer depth, strength of the thermocline, integral of fluorescence, to name just a few), as well as several metrics to describe the krill (number of aggregations, aggregation density, depth, and thickness). When I multiplied out how many possible combinations of predictor variables and parameters we’re interested in modeling, I realized this meant running nearly 300 models in order to settle on the best ten. This is where programming comes in, I told myself, and caught my breath.

I’ve always loved languages. When I was much younger, I thought I might want to study linguistics. As a graduate student in wildlife science, the language I’ve spent the most time learning, and come to love, is the statistical programming language R. Just like any other language, R has syntax and structure. Like any other language, there are many ways in which to articulate something, to make a particular point or reach a particular end goal. Well-written code is sometimes described as “elegant”, much like a well-articulated piece of writing. While I certainly do not consider myself “fluent” in R, it is a language I love learning. I like to think that the R scripts I write are an attempt to eloquently uncover and describe ecological patterns.

Rather than running 300 models one by one, I wrote an R script to run many models at a time, and then sort the outputs by model performance. I may look at the five best models of 32 options in order to select one. But this is where Leigh reminds me to step back from the programming for a minute and put my ecologist hat back on. Insight on the part of the modeler is needed in order to discern between what are real ecological relationships and what are spurious correlations in the data. It may not be quite as simple as choosing the model with the highest explanatory power when my goal is to make ecological inferences.

So, where does this leave me? Hundreds of models later, I am still not entirely sure which ones are best, although I’ve narrowed it down considerably. My programming proficiency and confidence continue to grow, but that only goes so far in ecology. Knowledge of my study system is equally important. So my workflow lately goes something like this: write code, try to interpret model outputs, consider what I know about the oceanography of my study region, re-write code, re-interpret the revised results, and so on. Hopefully this iterative process is bringing us gradually closer to an understanding of the ecology of blue whales on a foraging ground… stay tuned.

## Albatrosses at sunrise, dolphins at sunset: Northern California Current cruise

By Dawn Barlow, PhD student, Geospatial Ecology of Marine Megafauna Lab, Department of Fisheries and Wildlife, Oregon State University

Sun on my face and wind in my hair, scanning the expanse of blue. Forty minutes on, twenty minutes off, from sunrise until sunset, day after day. Hours of seemingly empty blue, punctuated by graceful black-footed albatrosses wheeling and gliding over the swells, by the splashing approach of a curious group of Pacific white-sided dolphins coming to play in the bow of the ship, by whale spouts on the horizon and the occasional breaching humpback. A flurry of data entry—geographic coordinates, bearing and distance from the ship, number of animals, species identification, behavior—and then back to blue.

I’ve just returned from the Northern California Current (NCC) ecosystem cruise aboard NOAA ship Bell M. Shimada. My role on board was the marine mammal observer, logging marine mammal sightings during the transits between sampling stations. We surveyed and sampled between Cape Mears, Oregon and Trinidad, California, from right along the coast out to 200 nautical miles offshore. Resources in the marine environment are patchy, and our coastline is highly productive. This diversity in environmental conditions creates niche habitats for many species, which is one reason why surveying and sampling across a broad geographic range can be so informative. We left Newport surrounded by gray whales, feeding in green, chilly waters at temperatures around 12°C. Moving west, the marine mammal and seabird sightings were increasingly sparse, the water increasingly blue, and the surface temperature warmed to a balmy 17°C. We had reached offshore waters, an ocean region sometimes referred to as the “blue desert”. For an entire day I didn’t see a single marine mammal and only just a few seabirds, until a handful of common dolphins—more frequently seen in warm-temperate and tropical waters to the south—joined the ship at sunset. As we transited back inshore over the productive Heceta Bank, the water became cooler and greener. I stayed busy logging sightings of humpback and gray whales, harbor porpoise and Dall’s porpoise, pacific white-sided dolphins and sea lions. These far-ranging marine predators must find a way to make a living in the patchy and dynamic ocean environment, and therefore their distribution is also patchy—aggregated around areas of high productivity and prey availability, and occasionally seen transiting in between.

Here are a few cruise highlights:

Curious groups of common dolphins (Delphinus delphis) came to play in the bow wake of the ship and even checked out the plankton nets when they were deployed. Common dolphins are typically found further south, however we saw several groups of them in the warmer waters far offshore.

Ocean sunfish (Mola mola) will occasionally lay themselves flat at the surface so that seabirds will pick them clean of any parasites. I was delighted to observe this for the first time just off Newport! There were several more sunfish sightings throughout the cruise.

A masked booby (Sula dactylatra) hung around the ship for a bit, 16 nautical miles from shore, just south of the Oregon-California border. Considered a tropical species, a sighting this far north is extremely rare. While masked boobies are typically distributed in the Caribbean and tropical Pacific from Mexico to Australia, one found its way to the Columbia River in 2006 (first record in the state of Oregon) and another showed up here to Newport in 2015 – reportedly only the second to be recorded north of Mendocino County, California. Perhaps this sighting is the third?

Masked booby (Sula dactylatra). Photo: Dawn Barlow.

While most of my boat-based fieldwork experiences have been focused on marine mammal research, this was an interdisciplinary cruise aimed at studying multiple aspects of the northern California current ecosystem. There were researchers on board studying oceanography, phytoplankton and harmful algal blooms, zooplankton, and microplastics. When a group of enthusiastic scientists with different areas of expertise come together and spend long days at sea, there is a wonderful opportunity to learn from one another. The hydroacoustic backscatter on the scientific echosounder prompted a group discussion about vertical migration of plankton one evening. Another evening I learned about differences in energetic content between krill species, and together we mused about what that might mean for marine predators. This is how collaborations are born, and I am grateful for the scientific musings with so many insightful people.

Thank you to the Shimada crew and the NCC science team for a wonderful cruise!

The NCC science team after a successful cruise!

## Cloudy with a chance of blue whales

By Dawn Barlow, PhD student, Department of Fisheries & Wildlife, Geospatial Ecology of Marine Megafauna Lab

As a PhD student studying the ecology of blue whales in New Zealand, my time is occupied by questions such as: When and where are the blue whales? Can we predict where they will be based on environmental conditions? How does their distribution overlap with human activity such as oil and gas exploration?

Leigh and I have just returned from New Zealand, where I gave an oral presentation at the Society for Conservation Biology Oceania Congress entitled “Cloudy with a chance of whales: Forecasting blue whale presence to mitigate industrial impacts based on tiered, bottom-up models”. While the findings I presented are preliminary, an exciting ecological story is emerging, and one with clear management implications.

The South Taranaki Bight (STB) region of New Zealand is an important area for a population of blue whales which are unique to New Zealand. A wind-driven upwelling system brings cold, productive waters into the bight [1], which sustains high densities of krill [2], blue whale prey. The region is also frequented by busy shipping traffic, oil and gas drilling and extraction platforms as well as seismic survey effort for subsurface oil and gas reserves, and is the site of a recently-permitted seabed mine for iron sands (Fig. 1). However, a lack of knowledge on blue whale distribution and habitat use patterns has impeded effective management of these potential anthropogenic threats.

Three surveys were conducted in the STB region in the summer months of 2014, 2016, and 2017. During that time, we not only looked for blue whales, we also collected oceanographic data and hydroacoustic backscatter data to map and measure aspects of the krill in the region. These data streams will help us understand the functional, ecological relationships between the environment (oceanography), prey (krill), and predators (blue whales) in the ecosystem (Fig. 2). But in practice these data are costly and time-consuming to collect, while other data sources such as satellite imagery are readily accessible to managers at a variety of spatial and temporal scales. Therefore, another one of my aims is to link the data we collected in the field to satellite imagery, so that managers can have a practical tool to predict when and where the blue whales are most likely to be found in the region.

So what did I find? Here are the highlights from my preliminary analyses:

• The majority of the patterns in blue whale distribution can be explained by the density, depth, and thickness of the krill patches.
• Patterns in the krill are driven by oceanography.
• Those same oceanographic parameters that drive the krill can be used to explain blue whale distribution.
• There are tight relationships between the important oceanographic variables and satellite images of sea surface temperature.
• Blue whale distribution can, to some degree, be explained using just satellite imagery.

We were able to identify a sea surface temperature range in the satellite imagery of approximately 18°C where the likelihood of finding a blue whale is the highest. Is this because blue whales really like 18° water? Well, more likely this relationship exists because the satellite imagery is reflective of the oceanography, and the oceanography drives patterns in the krill distribution, and the krill drives the distribution of blue whales (Fig. 3). We were able to make each of these functional linkages through our series of models, which is quite exciting.

That’s all well and good, but we were interested in testing these relationships to see if our identified habitat associations hold up even when we do not have field data (oceanographic, krill, and whale data). This past austral summer, we did not have a field season to collect data, but there was a large seismic airgun survey of the STB region. Seismic survey vessels are required to have trained marine mammal observers on board, and we were given access to the blue whale sightings data they recorded during the survey. In December, when the water was right around the preferred temperature identified by our models (18°C), the observers made 52 blue whale sightings (Fig. 4). In January and February, the waters warmed and only two sightings were made in each month. This is not only reassuring because it supports our model results, it also implies that there is the potential to balance industrial use of the area with protection of blue whale habitat, based on our understanding of the ecology. In January and February, very few blue whales were likely disturbed by the industrial activity in the STB, as conditions were not favorable for foraging at the location of the seismic survey. In contrast, the blue whales that were in the STB region in December may have experienced physiological consequences of sustained exposure to airgun noise since the conditions were favorable for foraging in the STB. In other words, the whales may have tolerated the noise exposure to gain access to good food, but this could have significant biological repercussions such as increased stress [3].

In the first two weeks of July, we presented these latest findings to managers at the New Zealand Department of Conservation, the Minister of Conservation, the CEO and Policy Advisor of a major oil and gas conglomerate, NGOs, advocacy groups, and scientific colleagues. It was valuable to gather feedback from many different stakeholders, and satisfying to see such a clear interest in, and management application of, our work.

What’s next? We’re back in Oregon, and diving back into analysis. We intend to take the modeling work a step further to make the models predictive—for example, can we forecast where the blue whales will be based on the temperature, productivity, and winds two weeks prior? I am excited to see where these next steps lead!

References:

1. Shirtcliffe TGL, Moore MI, Cole AG, Viner AB, Baldwin R, Chapman B. 1990 Dynamics of the Cape Farewell upwelling plume, New Zealand. New Zeal. J. Mar. Freshw. Res. 24, 555–568. (doi:10.1080/00288330.1990.9516446)
2. Bradford-Grieve JM, Murdoch RC, Chapman BE. 1993 Composition of macrozooplankton assemblages associated with the formation and decay of pulses within an upwelling plume in greater cook strait, New Zealand. New Zeal. J. Mar. Freshw. Res. 27, 1–22. (doi:10.1080/00288330.1993.9516541)
3. Rolland RM, Parks SE, Hunt KE, Castellote M, Corkeron PJ, Nowacek DP, Wasser SK, Kraus SD. 2012 Evidence that ship noise increases stress in right whales. Proc. Biol. Sci. 279, 2363–8. (doi:10.1098/rspb.2011.2429)

## Oregon sea otter reintroduction: opinions, perspectives, and theories

By Dominique Kone, Masters Student in Marine Resource Management

Species reintroductions can be hotly contested issues because they can negatively impact other species, ecosystems, and society, as well as failing, altogether. The uncertainty of their outcomes forces stakeholder groups to form their own opinions on whether it’s a good idea to proceed with a reintroduction. When you have several groups with conflicting values and views, managers need to focus on the information most important for them to make a well-informed decision on whether to pursue a reintroduction.

As researchers, we can play an important role by carefully considering and addressing these views through our research, if the appropriate data is available. Despite being in the early days of our study on the potential sea otter reintroduction to Oregon, we have already heard several perspectives regarding its potential success, the type of research we should do, and if sea otters should be brought back to Oregon. Here, I present some of the most interesting and relevant opinions, perspectives, and theories I’ve heard regarding this reintroduction idea.

The first reintroduction failed because of X, Y, and Z.

From 1970-1971, managers translocated 93 sea otters to Oregon in a reintroduction effort (Jameson et al. 1982). However, in a matter of 5-6 years, all sea otters disappeared, and the effort was considered a failure. Researchers have theorized that sea otters left Oregon due to a lack of suitable habitat and prey, or to return home to sites from which they were captured. Others have reasoned that managers should have introduced southern sea otters instead of northern sea otters, suggesting one subspecies’ genetic pre-disposition may improve their chance for survival.

Knowing the reasons for this failure may help managers avoid these causes in a future reintroduction attempt and increase its chance of success. We, as scientists, can also gain insight from knowing these causes because this may help us better tailor our research to potentially investigate whether those causes still pose a threat to sea otters during a second attempt. Unfortunately, we lack concrete evidence on what exactly caused this failure, but we can still work to test some these theories.

An otter is an otter, no matter where you put it.

There is evidence that northern and southern sea otters are genetically distinct, to a certain degree (Valentine et al. 2008, Larson et al. 2012), and hypotheses have been put forward that the two subspecies may be behaviorally- and ecologically-distinct, too. Studies have shown that northern and southern sea otters have different sized and shaped skulls and teeth, which researchers hypothesize may be a specialized foraging adaptation for consuming different prey species (Campbell & Santana 2017, Timm-Davis et al. 2015). This view suggests that each subspecies has developed unique traits to adapt to the environmental conditions specific to their current ranges. Therefore, when considering which subspecies to bring to Oregon, managers should reintroduce the subspecies with traits better-suited to cope with the types of habitat, prey assemblages, and oceanographic conditions specific to Oregon.

However, other scientists hold the opposite view, and argue that “an otter is an otter” no matter where you put it. This perspective suggests that both subspecies have an equal chance at surviving in any type of suitable habitat because all otters behave in similar ways. Therefore, ecologically, it may not matter which subspecies managers bring to Oregon.

Oregon doesn’t have enough sea otter habitat.

Kelp is considered important sea otter habitat. In areas with high sea otter densities, such as central and southern California, kelp forests are persistent throughout the year. However, in Oregon, our kelp primarily consists of bull kelp – a slightly more fragile species compared to the durable giant kelp in California. In winter, this bull kelp gets dislodged during intense storms, resulting in seasonal changes in kelp availability. Managers worry that this seasonality could reduce the amount of suitable habitat, to the point where Oregon may not be able to support sea otters.

Yet, we know sea otters used to exist here; therefore, we can assume there must have been some suitable habitat that may persist today. Furthermore, sea otters use a range of habitats, including estuaries, bays, and reefs (Laidre et al. 2009, Lafferty & Tinker 2014, Kvitek et al. 1988). Therefore, even during times when kelp is less abundant, sea otters could use these other forms of habitat along the Oregon coast. Luckily, we have the spatial tools and data to assess how much, where, and when we have suitable habitat, and I will specifically address this in my thesis.

They’ll eat everything!

Sea otters are famous for their voracious appetites for benthic invertebrates, some of which are of commercial and recreational importance to nearshore fisheries. In some cases, sea otters have significantly reduced prey densities, such as sea urchins and Dungeness crab (Garshelis & Garshelis 1984, Estes & Palmisano 1974). However, without a formal analysis, it’s difficult to know if sea otters will have similar impacts on Oregon’s nearshore species, as well as at spatial scale these impacts will occur and whether our fisheries will be affected. We can predict where sea otters are likely to occur based on the presence of suitable habitat, but foraging impacts could be more localized or widespread across sea otter’s entire potential range. To better anticipate these impacts, managers will need an understanding of how much sea otters eat, where foraging could occur based on the availability of prey, and where sea otters and fisheries are likely to interact. I will also address this concern in my thesis.

To reintroduce or not to reintroduce? That is the question.

I have found that many scientists and managers have strong opinions on whether it’s appropriate to bring sea otters back to Oregon. Those who argue against a reintroduction often highlight many of the theories already mentioned here – lack of habitat, potential impacts to fisheries, and genetics. While other opponents provided more logistical and practical justifications, such as confounding politics, as well as difficulties in getting public support and regulatory permission to move a federally-listed species.

In contrast, proponents of this idea argue that a reintroduction could augment the recovery of the species by providing additional habitat for the species to rebound to pre-exploitation levels, as well as allowing for increased gene flow between southern and northern sea otter populations. Other proponents have brought up potential benefits to humans, such restoring ecosystem services, providing an economic boost through tourism, or preserving tribal and cultural connections. Such benefits may be worth attempting another reintroduction effort.

As you can see, there are several opinions and perspectives related to a potential sea otter reintroduction to Oregon. While it’s important to consider all opinions, managers still need facts to make key decisions. Scientists can play an important role in providing this information, so managers can make a well-informed decision. Oregon managers have not yet decided whether to proceed with a sea otter reintroduction, but our lab is working to provide them with reliable and accurate science, so they may form their own opinions and arrive at their own decision.

References:

Estes, J. A. and J. F. Palmisano. 1974. Sea otters: the role in structuring nearshore communities. Science. 185: 1058-1060.

Garshelis, D. L. and J. A. Garshelis. 1984. Movements and management of sea otters in Alaska. The Journal of Wildlife Management. 48: 665-678.

Jameson, R. J, Kenyon, K. W., Johnson, A. M., and H. M. Wight. 1982. History and status of translocated sea otter populations in North America. Wildlife Society Bulletin. 10: 100-107.

Lafferty, K. D., and M. T. Tinker. 2014. Sea otters are recolonizing southern California in fits and starts. Ecosphere. 5(5).

Laidre, K. L., Jameson, R. J., Gurarie, E., Jeffries, S. J., and H. Allen. 2009. Spatial habitat use patterns of sea otters in coastal Washington. Journal of Mammalogy. 90(4): 906-917.

Kvitek, R. G. ,Fukayama, A. K., Anderson, B. S., and B. K. Grimm. 1988. Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon. Marine Biology. 98: 157-167.

Larson, S., Jameson, R., Etnier, M., Jones, T., and R. Hall. 2012. Genetic diversity and population parameters of sea otters, Enhydra lutris, before fur trade extirpation from 1741-1911. PLoS ONE. 7(3).

Timm-Davis, L. L, DeWitt, T. J., and C. D. Marshall. 2015. Divergent skull morphology supports two trophic specializations in otters (Lutrinae). PLoS ONE. 10(12).

Valentine et al. 2008. Ancient DNA reveals genotypic relationships among Oregon populations of the sea otter (Enhydra lutris). Conservation Genetics. 9:933-938.

## Forecasting blue whale presence: Small steps toward big goals

By Dawn Barlow, MSc student, Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

In 2013, Leigh first published a hypothesis that the South Taranaki Bight region between New Zealand’s North and South Islands is important habitat for blue whales  (Torres 2013). Since then, we have collected three years of data and conducted dedicated analyses, so we now understand that a unique population of blue whales is found in New Zealand, and that they are present in the South Taranaki Bight year-round (Barlow et al. in press).

This research has garnered quite a bit of political and media attention. A major platform item for the New Zealand Green Party around the last election was the establishment of a marine mammal sanctuary in the South Taranaki Bight. When the world’s largest seismic survey vessel began surveying the South Taranaki Bight this summer for more oil and gas reserves using tremendously loud airguns, there were rallies on the lawn in front of Parliament featuring a large inflatable blue whale that the protesters affectionately refer to as “Janet”. Needless to say, blue whales have made their way into the spotlight in New Zealand.

Now that we know there is a unique population of blue whales in New Zealand, what is next? What’s next for me is an exciting combination of both ecology and conservation. If an effective sanctuary is to be implemented, it needs to be more than a simple box drawn on a map to check off a political agenda item—the sanctuary should be informed by our best ecological knowledge of the blue whales and their habitat.

In July, Leigh and I will attend the Society for Conservation Biology meeting in Wellington, New Zealand, and I’ll be giving a presentation titled “Cloudy with a chance of whales: Forecasting blue whale presence based on tiered, bottom-up models”. I’ll be the first to admit, I am not yet forecasting blue whale presence. But I am working my way there, step-by-step, through this tiered, bottom-up approach. In cetacean habitat modeling, we often assume that whale distribution on a foraging ground is determined by their prey’s distribution, and that satellite images of temperature and chlorophyll-a provide an accurate picture of what is going on below the surface. Is this true? With our three years of data including in situ oceanography, krill hydroacoustics, and blue whale distribution and behavior, we are in a unique position to test some of those assumptions, as well as provide managers with an informed management tool to predict blue whale distribution.

What questions will we ask using our data? Firstly, can in situ oceanography (i.e., thermocline depth and temperature, mixed layer depth) predict the distribution and density of blue whale prey (krill)? Then, can those prey patterns be accurately predicted in the absence of oceanographic measurements, using just satellite images? Next, we’ll bring the blue whales back into the picture to ask: can we predict blue whale distribution based on our in situ measurements of oceanography and prey? And finally, in the absence of in situ measurements (which is most often the case), can we forecast where the whales will be based just on remotely-sensed images of the region?

So, cloudy with a chance of whales? Well, you’ll have to stay tuned for that story in the coming months. In the meantime, I can tell you that as daunting as it is to aggregate so many data streams, each step of the way has a piece of the story to tell. I can’t wait to see how it falls together, both from an ecological modeling perspective and a conservation management objective.

References:

Torres, L. G. (2013). Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zealand Journal of Marine and Freshwater Research47(2), 235-248.

Barlow, D. R., Torres, L. G., Hodge, K. B., Steel, D. Baker, C. S., Chandler, T. E., Bott, N., Constantine, R., Double, M. C., Gill, P., Glasgow, D., Hamner, R. M., Lilley, C., Ogle, M., Olson, P. A., Peters, C., Stockin, K. A., Tessaglia-Hymes, C. T., Klinck, H. (in press). Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endangered Species Research.

## The Land of Maps and Charts: Geospatial Ecology

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

I love maps. I love charts. As a random bit of trivia, there is a difference between a map and a chart. A map is a visual representation of land that may include details like topology, whereas a chart refers to nautical information such as water depth, shoreline, tides, and obstructions.

I have an intense affinity for visually displaying information. As a child, my dad traveled constantly, from Barrow, Alaska to Istanbul, Turkey. Immediately upon his return, I would grab our standing globe from the dining room and our stack of atlases from the coffee table. I would sit at the kitchen table, enthralled at the stories of his travels. Yet, a story was only great when I could picture it for myself. (I should remind you, this was the early 1990s, GoogleMaps wasn’t a thing.) Our kitchen table transformed into a scene from Master and Commander—except, instead of nautical charts and compasses, we had an atlas the size of an overgrown toddler and salt and pepper shakers to pinpoint locations. I now had the world at my fingertips. My dad would show me the paths he took from our home to his various destinations and tell me about the topography, the demographics, the population, the terrain type—all attribute features that could be included in common-day geographic information systems (GIS).

As I got older, the kitchen table slowly began to resemble what I imagine the set from Master and Commander actually looked like; nautical charts, tide tables, and wind predictions were piled high and the salt and pepper shakers were replaced with pencil marks indicating potential routes for us to travel via sailboat. The two of us were in our element. Surrounded by visual and graphical representations of geographic and spatial information: maps. To put my map-attraction this in even more context, this is a scientist who grew up playing “Take-Off”, a board game that was “designed to teach geography” and involved flying your fleet of planes across a Mercator projection-style mapboard. Now, it’s no wonder that I’m a graduate student in a lab that focuses on the geospatial aspects of ecology.

So why and how did geospatial ecology became a field—and a predominant one at that? It wasn’t that one day a lightbulb went off and a statistician decided to draw out the results. It was a progression, built upon for thousands of years. There are maps dating back to 2300 B.C. on Babylonian clay tablets (The British Museum), and yet, some of the maps we make today require highly sophisticated technology. Geospatial analysis is dynamic. It’s evolving. Today I’m using ArcGIS software to interpolate mass amounts of publicly-available sea surface temperature satellite data from 1981-2015, which I will overlay with a layer of bottlenose dolphin sightings during the same time period for comparison. Tomorrow, there might be a new version of software that allows me to animate these data. Heck, it might already exist and I’m not aware of it. This growth is the beauty of this field. Geospatial ecology is made for us cartophiles (map-lovers) who study the interdependency of biological systems where location and distance between things matters.

In a broader context, geospatial ecology communicates our science to all of you. If I posted a bunch of statistical outputs in text or even table form, your eyes might glaze over…and so might mine. But, if I displayed that same underlying data and results on a beautiful map with color-coded symbology, a legend, a compass rose, and a scale bar, you might have this great “ah-ha!” moment. That is my goal. That is what geospatial ecology is to me. It’s a way to SHOW my science, rather than TELL it.

Would you like to see this over and over again…?

Or see this once…?

For many, maps are visually easy to interpret, allowing quick message communication. Yet, there are many different learning styles. From my personal story, I think it’s relatively obvious that I’m, at least partially, a visual learner. When I was in primary school, I would read the directions thoroughly, but only truly absorb the material once the teacher showed me an example. Set up an experiment? Sure, I’ll read the lab report, but I’m going to refer to the diagrams of the set-up constantly. To this day, I always ask for an example. Teach me a new game? Let’s play the first round and then I’ll pick it up. It’s how I learned to sail. My dad described every part of the sailboat in detail and all I heard was words. Then, my dad showed me how to sail, and it came naturally. It’s only as an adult that I know what “that blue line thingy” is called. Geospatial ecology is how I SEE my research. It makes sense to me. And, hopefully, it makes sense to some of you!

I strongly believe a meaningful career allows you to highlight your passions and personal strengths. For me, that means photography, all things nautical, the great outdoors, wildlife conservation, and maps/charts.  If I converted that into an equation, I think this is a likely result:

Photography + Nautical + Outdoors + Wildlife Conservation + Maps/Charts = Geospatial Ecology of Marine Megafauna

Or, better yet:

? + ⚓ + ? + ? + ? =  GEMM Lab

This lab was my solution all along. As part of my research on common bottlenose dolphins, I work on a small inflatable boat off the coast of California (nautical ✅, outdoors ✅), photograph their dorsal fin (photography ✅), and communicate my data using informative maps that will hopefully bring positive change to the marine environment (maps/charts ✅, wildlife conservation✅). Geospatial ecology allows me to participate in research that I deeply enjoy and hopefully, will make the world a little bit of a better place. Oh, and make maps.