Inference, and the intersection of ecology and statistics

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

Recently, I had the opportunity to attend the International Statistical Ecology Conference (ISEC), a biennial meeting of researchers at the interface of ecology and statistics. I am a marine ecologist, fascinated by the interactions between animals and the dynamic ocean environment they inhabit. If you had asked me five years ago whether I thought I would ever consider myself a statistician or a computer programmer, my answer would certainly have been “no”. Now, I find myself studying the ecology of blue whales in New Zealand using a variety of data streams and methodologies, but a central theme for my dissertation is species distribution modeling. Species distribution models (SDMs) are mathematical algorithms that correlate observations of a species with environmental conditions at their observed locations to gain ecological insight and predict spatial distributions of the species (Fig. 1; Elith and Leathwick 2009). I still can’t say I would identify as a statistician, but I have a growing appreciation for the role of statistics to gain inference in ecology.

Figure 1. A schematic of a species distribution model (SDM) illustrating how the relationship between mapped species and environmental data (left) is compared to describe “environmental space” (center), and then map predictions from a model using only environmental predictors (right). Note that inter-site distances in geographic space might be quite different from those in environmental space—a and c are close geographically, but not environmentally. The patterning in the predictions reflects the spatial autocorrelation of the environmental predictors. Figure reproduced from Elith and Leathwick (2009).

Before I continue, let’s take a look at just a few definitions from Merriam-Webster’s dictionary:

Statistics: a branch of mathematics dealing with the collection, analysis, interpretation, and presentation of masses of numerical data

Ecology: a branch of science concerned with the interrelationship of organisms and their environments

Inference: a conclusion or opinion that is formed because of known facts or evidence

Ecological data are notoriously noisy, messy, and complex. Statistical tests are meant to help us understand whether a pattern in the data is different from what we would expect through random chance. When we study how organisms interact with one another and their environment, it is impossible to completely capture all elements of the ecosystem. Therefore, ecology is a field ripe with challenges for statisticians. How do we quantify a meaningful biological signal amidst all the noise? How can we gain inference from ecological data to enhance knowledge, and how can we use that knowledge to make informed predictions? Marine mammals are notoriously difficult to study. They inhabit an environment that is relatively inaccessible and inhospitable to humans, they occur in low numbers, they are highly mobile, and they are rarely visible. All ecological data are difficult and noisy and riddled with small sample sizes, but counting trees presents fewer logistical challenges than counting moving whales in an ever-changing open-ocean setting. Therefore, new methodologies in areas like species distribution modeling are often developed using large, terrestrial datasets and eventually migrate to applications in the marine environment (Robinson et al. 2011).

Many presentations I attended at the conference were geared toward moving beyond correlative SDMs. SDMs were developed to correlate species occurrence patterns with features of the environment they inhabit (e.g. temperature, precipitation, terrain, etc.). However, those relationships do not actually explain the underlying mechanism of why a species is more likely to occur in one environment compared to another. Therefore, ecological statisticians are now using additional information and modeling approaches within SDMs to incorporate information such as species co-occurrence patterns, population demographic information, and physiological constraints. Building SDMs to include such process-explicit information allows us to make steps toward understanding not just when and where a species occurs, but why.

Machine learning is an area that continues to advance and open doors to new applications in ecology. Machine learning approaches differ fundamentally from classical statistics. In statistics, we formulate a hypothesis, select the appropriate model to test that hypothesis (for example, linear regression), then test how well the data fit the model (“Is the relationship linear?”), and test the strength of that inference (“Is the linear pattern different from what we would expect due to random chance?”). Machine learning, on the other hand, does not use a predetermined notion of relationships between variables. Rather, it tries to create an algorithm that fits the patterns in the data. Statistics asks how well the data fit a model, and machine learning asks how well a model fits the data.

Machine learning approaches allow for very complex relationships to be included in models and can be excellent for making predictions. However, sometimes the relationships fitted by a machine learning algorithm are so complex that it is not possible to infer any ecological meaning from them. As one ISEC presenter put it, in machine learning “the computer learns but the scientist does not”. The most important thing when selecting your methodology is to remember your question and your goal. Do you want to understand the mechanism of why an animal is where it is? Or do you not need to understand the driver, but rather want to make the best predictions of where an animal will be? In my case, the answer to that question differs from one of my PhD chapters to the next. We want to understand the functional relationships between oceanography, krill availability, and blue whale distribution (Barlow et al. 2020), and subsequently we want to develop forecasting models that can reliably predict blue whale distribution to inform conservation efforts (Fig. 2).

Figure 2. An example predictive map of where we expect blue whales to be distributed based on environmental conditions. Warmer colors represent areas with a higher probability of blue whale occurrence, and the blue crosses represent locations where blue whales were observed.

ISEC was an excellent opportunity for me to break out of my usual marine mammal-centered bubble and get a taste of what is happening on the leading edge of statistical ecology. I learned about the latest approaches and innovations in species distribution modeling, and in the process I also learned about trees, koalas, birds, and many other organisms from around the world. A fun bonus of attending a methods-focused conference is learning about completely new study species and systems. There are many ways of approaching an ecological question, gaining inference, and making predictions. I look forward to incorporating the knowledge I gained through ISEC into my own research, both in my doctoral work and in applications of new methods to future research projects.

Figure 3. The virtual conference photo of all who attended the biennial International Statistical Ecology Conference. Thank you to the organizers, who made it a truly excellent and engaging conference experience!

References

Barlow, D.R., Bernard, K.S., Escobar-Flores, P., Palacios, D.M., and 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. Mar. Ecol. Prog. Ser. doi:https://doi.org/10.3354/meps13339.

Elith, J., and Leathwick, J.R. 2009. Species Distribution Models: Ecological Explanation and Prediction Across Space and Time. Annu. Rev. Ecol. Evol. Syst. 40(1): 677–697. doi:10.1146/annurev.ecolsys.110308.120159.

Robinson, L.M., Elith, J., Hobday, A.J., Pearson, R.G., Kendall, B.E., Possingham, H.P., and Richardson, A.J. 2011. Pushing the limits in marine species distribution modelling: Lessons from the land present challenges and opportunities. doi:10.1111/j.1466-8238.2010.00636.x.

The teamwork of conservation science

Dr. Leigh Torres
PI, Geospatial Ecology of Marine Megafauna Lab, Marine Mammal Institute
Assistant Professor, Oregon Sea Grant, Department of Fisheries and Wildlife, Oregon State University

I have played on sports teams all my life – since I was four until present day. Mostly soccer teams, but a fair bit of Ultimate too. Teams are an interesting beast. They can be frustrating when communication breaks down, irritating when everyone is not on the same timeline, and disastrous if individuals do not complete their designated job. Yet, without the whole team we would never win. So, on top of the fun of competition, skill development, and exercise, playing on teams has always been part of the challenging and fulfilling process for me: everyone working toward the same goal – to win – by making the team fluid, complimentary, integrated, and ultimately successful.

I have come to learn that it is the same with conservation science.

A few of my teams through the ages, as player and coach. Some of my favorite people are on these teams, from 1981 to 2018.

Conservation efforts are often so complex, that it is practically impossible to achieve success alone. Forces driving the need for conservation typically include monetary needs/desires, social values, ecological processes, animal physiology, multi-jurisdictional policies, and human behavior. Each one of these forces alone is challenging to understand and takes expertise to comprehend the situation. Hence, building a well-functioning team is essential. Here’s a recent example from the GEMM Lab:

Since 2014 entanglements of blue, humpback and gray whales in fishing gear along the west coast of the USA have dramatically increased, particularly in Dungeness crab fishing gear. Many forces likely led to this increase, including increased whale population abundance, potential shifts in whale distributions, and changes in fishing fleet dynamics. While we cannot point a finger at one cause, many people and groups recognize that we cannot continue to let whales become entangled and killed at such high rates: whale populations would decline, fisheries would look bad in the public eye and potentially lose profits, whales have an intrinsic right to live in the ocean without being bycaught, and whales are an important part of the ecosystem that would deteriorate without them. In 2017, the Oregon Whale Entanglement Working Group was formed to bring stakeholders together that were concerned about this problem to discuss possible solutions and paths forward. I was lucky to be a part of this group, which also included members of the Dungeness crab fishery and commission, the Oregon Department of Fish and Wildlife (ODFW), other marine mammal scientists, and representatives of the American Cetacean Society, The Nature Conservancy, and a local marine gear supplier.

We met regularly over 2.5 years, and despite some hesitation at first about walking into a room of potentially disgruntled fishermen (I would be lying if I did not admit to this), after the first meeting I looked forward to every gathering. I learned an immense amount about the Dungeness crab fishery and how it operates, how ODFW manages the fishery and why, and what people do, don’t and need to know about whales in Oregon. Everyone agreed that reducing whale entanglements is needed, and a frequent approach discussed was to reduce risk by not setting gear where and when we expect whales to be. Yet, this idea flagged a very critical knowledge gap: We do not have a good understanding of whale distribution patterns in Oregon. Thus leading to the development of a highly collaborative research effort to describe whale distribution patterns in Oregon and identify areas of co-occurrence between whales and fishing effort to reduce the risk of entanglements. Sounds great, but a tough task to accomplish in a few short years. So, let me introduce the great team I am working with to make it all happen.

While I may know a few things about whales and spatial ecology, I don’t know too much about fisheries in Oregon. My collaboration with folks at ODFW, particularly Kelly Corbett and Troy Buell, has enabled this project to develop and go forward, and ultimately will lead to success. These partners provide feedback about how and where the fishery operates so I know where and when to collect data, and importantly they will provide the information on fishing effort in Oregon waters to relate to our generated maps of whale distribution. This spatial comparison will produce what is needed by managers and fishermen to make informed and effective decisions about where to fish, and not to fish, so that we reduce whale entanglement risk while still harvesting successfully to ensure the health and sustainability of our coastal economies.

So, how can we collect standardized data on whale distribution in Oregon waters without breaking the bank? I tossed this question around for a long time, and then I looked up to the sky and wondered what that US Coast Guard (USCG) helicopter was flying around for all the time. I reached out to the USCG to enquire, and proposed that we have an observer fly in the helicopter with them along a set trackline during their training flights. Turns out the USCG Sector North Bend and Columbia River were eager to work with us and support our research. They have turned out to be truly excellent partners in this work. We had some kinks to work out at the beginning – lots of acronyms, protocols, and logistics for both sides to figure out – but everyone has been supportive and pleasant to work with. The pilots and crew are interested in our work and it is a joy to hear their questions and see them learn about the marine ecosystem. And our knowledge of helicopter navigation and USCG duties has grown astronomically.

On the left is a plot of the four tracklines we survey for whales each month for two years aboard a US Coast Guard helicopter. On the right are some photos of us in action with our Coast Guard partners.

Despite significant cost savings to the project through our partnership with the USCG, we still need funds to support time, gear and more. And full credit to the Oregon Dungeness Crab Commission for recognizing the value and need for this project to support their industry, and stepping up to fund the first year of this project. Without their trust and support the project may not have got off the ground. With this support in our back pocket and proof of our capability, ODFW and I teamed up to approach the National Oceanographic and Atmospheric and Administration (NOAA) for funds to support the remaining years of the project. We found success through the NOAA Fisheries Endangered Species Act Section 6 Program, and we are now working toward providing the information needed to protect endangered and threatened whales in Oregon waters.

Despite our cost-effective and solid approach to data collection on whale occurrence, we cannot be everywhere all the time looking for whales. So we have also teamed up with Amanda Gladics at Oregon Sea Grant to help us with an important outreach and citizen science component of the project. With Amanda we have developed brochures and videos to inform mariners of all kinds about the project, objectives, and need for them to play a part. We are encouraging everyone to use the Whale Alert app to record their opportunistic sightings of whales in Oregon waters. These data will help us build and test our predictive models of whale distribution. Through this partnership we continue important conversations with fishermen from many fisheries about their concerns, where they are seeing whales, and what needs to be done to solve this complex conservation challenge.  

Of course I cannot collect, process, analyze, and interpret all this data on my own. I do not have the skills or capacity for that. My partner in the sky is Craig Hayslip, a Faculty Research Assistant in the Marine Mammal Institute. Craig has immense field experience collecting data on whales and is the primary observer on the survey flights. Together we have navigated the USCG world and developed methods to collect our data effectively and efficiently (all within a tiny space flying over the ocean). In a few months we will be ¾ of the way through our data collection phase, which means data analysis will take over. For this phase I am bringing back a GEMM Lab star, Solene Derville, who recently completed her PhD. As the post-doc on the project, Solene will take the lead on the species distribution modeling and fisheries overlap analysis. I am looking forward to partnering with Solene again to compile multiple data sources on whales and oceanography in Oregon to produce reliable and accurate predictions of whale occurrence and entanglement risk. Finally I want to acknowledge our great partners at the Cascadia Research Collective (Olympia, WA) and the Cetacean Conservation and Genomics Lab (OSU, Marine Mammal Institute) who help facilitate our data collection, and conduct the whale photo-identification or genetic analyses to determine population assignment.  

As you can see, even this one, smallish, conservation research project takes a diverse team of partners to proceed and ensure success. On this team, my position is sometimes a player, coach, or manager, but I am always grateful for these amazing collaborations and opportunities to learn. I am confident in our success and will report back on our accomplishments as we wrap up this important and exciting conservation science project.   

A fin whale observed off the Oregon coast during one of our surveys aboard a US Coast Guard helicopter.

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.

A pair of blue whales surfacing in the South Taranaki Bight region of New Zealand. Drone piloted by Todd Chandler during the 2017 field season.

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.

A blue whale breaks the surface. As I grapple with questions of model complexity and scale of analysis, I sometimes need a reminder that behind each data point is a blue whale, and what a privilege it is to study them. Photo by Leigh Torres.

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.

The study species: common bottlenose dolphin (Tursiops truncatus) seen along the California coastline in 2015. Image source: Alexa Kownacki.

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.

Figure 1. Map representing the San Diego transect, represented with a light blue line inside of a one-kilometer buffered “sighting zone” in pale yellow. The dark pink shapes are dolphin sightings from 1981-2015, although some are stacked on each other and cannot be differentiated. The lagoons, ranging in size, are color-coded. The transect line runs from the breakwaters of Mission Bay, CA to Oceanside Harbor, CA.

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.

Figure 2. Summary from the Average Nearest Neighbor calculation in ArcMap 10.6 displaying that bottlenose dolphin sightings in San Diego are highly clustered. When the z-score, which corresponds to different colors on the graphic above, is strongly negative (< -2.58), in this case dark blue, it indicates clustering. Because the p-value is very small, in this case, much less than 0.01, these results of clustering are strongly significant.

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.

Figure 3. Histogram comparing the latitudes with the frequency of dolphin sightings in San Diego, CA. The x-axis represents the latitudinal difference from the most northern part of the transect to each dolphin sighting. Therefore, a small difference would translate to a sighting being in the northern transect areas whereas large differences would translate to sightings being more southerly. This could be read from left to right as most northern to most southern. The y-axis represents the frequency of which those differences are seen, that is, the number of sightings with that amount of latitudinal difference, or essentially location on the transect line. Therefore, you can see there is a peak in the number of sightings towards the southern part of the transect line.

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.

Figure 4. Boxplot visualizing distributions of dolphin sightings latitudinal differences and ENSO index, with -1,0, and 1 representing cold, neutral, and warm years, respectively.

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.

Figure 5. Bar plot displaying the different distances from dolphin sighting location to the nearest lagoon in San Diego in decimal degrees. Note: Lagoon 4 is south of the study site and therefore was never the nearest 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.

Figure 6. Violin plot illustrating the distance from lagoons of dolphin sightings under different ENSO conditions. There are three major groups based on ENSO index: “-1” representing cold years, “0” representing neutral years, and “1” representing warm years. On the x-axis are lagoon IDs and on the y-axis is the distance to the nearest lagoon in decimal degrees. The wider the shapes, the more sightings, therefore Lagoon 6 has many sightings within a very small distance compared to Lagoon 5 where sightings are widely dispersed at greater distances.

 

Bottlenose dolphins foraging in a small group along the California coast in 2015. Image source: Alexa Kownacki.

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.

Data Wrangling to Assess Data Availability: A Data Detective at Work

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

Data wrangling, in my own loose definition, is the necessary combination of both data selection and data collection. Wrangling your data requires accessing then assessing your data. Data collection is just what it sounds like: gathering all data points necessary for your project. Data selection is the process of cleaning and trimming data for final analyses; it is a whole new bag of worms that requires decision-making and critical thinking. During this process of data wrangling, I discovered there are two major avenues to obtain data: 1) you collect it, which frequently requires an exorbitant amount of time in the field, in the lab, and/or behind a computer, or 2) other people have already collected it, and through collaboration you put it to a good use (often a different use then its initial intent). The latter approach may result in the collection of so much data that you must decide which data should be included to answer your hypotheses. This process of data wrangling is the hurdle I am facing at this moment. I feel like I am a data detective.

Data wrangling illustrated by members of the R-programming community. (Image source: R-bloggers.com)

My project focuses on assessing the health conditions of the two ecotypes of bottlenose dolphins between the waters off of Ensenada, Baja California, Mexico to San Francisco, California, USA between 1981-2015. During the government shutdown, much of my data was inaccessible, seeing as it was in possession of my collaborators at federal agencies. However, now that the shutdown is over, my data is flowing in, and my questions are piling up. I can now begin to look at where these animals have been sighted over the past decades, which ecotypes have higher contaminant levels in their blubber, which animals have higher stress levels and if these are related to geospatial location, where animals are more susceptible to human disturbance, if sex plays a role in stress or contaminant load levels, which environmental variables influence stress levels and contaminant levels, and more!

Alexa, alongside collaborators, photographing transiting bottlenose dolphins along the coastline near Santa Barbara, CA in 2015 as part of the data collection process. (Image source: Nick Kellar).

Over the last two weeks, I was emailed three separate Excel spreadsheets representing three datasets, that contain partially overlapping data. If Microsoft Access is foreign to you, I would compare this dilemma to a very confusing exam question of “matching the word with the definition”, except with the words being in different languages from the definitions. If you have used Microsoft Access databases, you probably know the system of querying and matching data in different databases. Well, imagine trying to do this with Excel spreadsheets because the databases are not linked. Now you can see why I need to take a data management course and start using platforms other than Excel to manage my data.

A visual interpretation of trying to combine datasets being like matching the English definition to the Spanish translation. (Image source: Enchanted Learning)

In the first dataset, there are 6,136 sightings of Common bottlenose dolphins (Tursiops truncatus) documented in my study area. Some years have no sightings, some years have fewer than 100 sightings, and other years have over 500 sightings. In another dataset, there are 398 bottlenose dolphin biopsy samples collected between the years of 1992-2016 in a genetics database that can provide the sex of the animal. The final dataset contains records of 774 bottlenose dolphin biopsy samples collected between 1993-2018 that could be tested for hormone and/or contaminant levels. Some of these samples have identification numbers that can be matched to the other dataset. Within these cross-reference matches there are conflicting data in terms of amount of tissue remaining for analyses. Sorting these conflicts out will involve more digging from my end and additional communication with collaborators: data wrangling at its best. Circling back to what I mentioned in the beginning of this post, this data was collected by other people over decades and the collection methods were not standardized for my project. I benefit from years of data collection by other scientists and I am grateful for all of their hard work. However, now my hard work begins.

The cutest part of data wrangling: finding adorable images of bottlenose dolphins, photographed during a coastal survey. (Image source: Alexa Kownacki).

There is also a large amount of data that I downloaded from federally-maintained websites. For example, dolphin sighting data from research cruises are available for public access from the OBIS (Ocean Biogeographic Information System) Sea Map website. It boasts 5,927,551 records from 1,096 data sets containing information on 711 species with the help of 410 collaborators. This website is incredible as it allows you to search through different data criteria and then download the data in a variety of formats and contains an interactive map of the data. You can explore this at your leisure, but I want to point out the sheer amount of data. In my case, the OBIS Sea Map website is only one major platform that contains many sources of data that has already been collected, not specifically for me or my project, but will be utilized. As a follow-up to using data collected by other scientists, it is critical to give credit where credit is due. One of the benefits of using this website, is there is information about how to properly credit the collaborators when downloading data. See below for an example:

Example citation for a dataset (Dataset ID: 1201):

Lockhart, G.G., DiGiovanni Jr., R.A., DePerte, A.M. 2014. Virginia and Maryland Sea Turtle Research and Conservation Initiative Aerial Survey Sightings, May 2011 through July 2013. Downloaded from OBIS-SEAMAP (http://seamap.env.duke.edu/dataset/1201) on xxxx-xx-xx.

Citation for OBIS-SEAMAP:

Halpin, P.N., A.J. Read, E. Fujioka, B.D. Best, B. Donnelly, L.J. Hazen, C. Kot, K. Urian, E. LaBrecque, A. Dimatteo, J. Cleary, C. Good, L.B. Crowder, and K.D. Hyrenbach. 2009. OBIS-SEAMAP: The world data center for marine mammal, sea bird, and sea turtle distributions. Oceanography 22(2):104-115

Another federally-maintained data source that boasts more data than I can quantify is the well-known ERDDAP website. After a few Google searches, I finally discovered that the acronym stands for Environmental Research Division’s Data Access Program. Essentially, this the holy grail of environmental data for marine scientists. I have downloaded so much data from this website that Excel cannot open the csv files. Here is yet another reason why young scientists, like myself, need to transition out of using Excel and into data management systems that are developed to handle large-scale datasets. Everything from daily sea surface temperatures collected on every, one-degree of latitude and longitude line from 1981-2015 over my entire study site to Ekman transport levels taken every six hours on every longitudinal degree line over my study area. I will add some environmental variables in species distribution models to see which account for the largest amount of variability in my data. The next step in data selection begins with statistics. It is important to find if there are highly correlated environmental factors prior to modeling data. Learn more about fitting cetacean data to models here.

The ERDAPP website combined all of the average Sea Surface Temperatures collected daily from 1981-2018 over my study site into a graphical display of monthly composites. (Image Source: ERDDAP)

As you can imagine, this amount of data from many sources and collaborators is equal parts daunting and exhilarating. Before I even begin the process of determining the spatial and temporal spread of dolphin sightings data, I have to identify which data points have sex identified from either hormone levels or genetics, which data points have contaminants levels already quantified, which samples still have tissue available for additional testing, and so on. Once I have cleaned up the datasets, I will import the data into the R programming package. Then I can visualize my data in plots, charts, and graphs; this will help me identify outliers and potential challenges with my data, and, hopefully, start to see answers to my focal questions. Only then, can I dive into the deep and exciting waters of species distribution modeling and more advanced statistical analyses. This is data wrangling and I am the data detective.

What people may think a ‘data detective’ looks like, when, in reality, it is a person sitting at a computer. (Image source: Elder Research)

Like the well-known phrase, “With great power comes great responsibility”, I believe that with great data, comes great responsibility, because data is power. It is up to me as the scientist to decide which data is most powerful at answering my questions.

Data is information. Information is knowledge. Knowledge is power. (Image source: thedatachick.com)

 

Finding the right fit: a journey into cetacean distribution models

Solène Derville, Entropie Lab, French National Institute for Sustainable Development (IRD – UMR Entropie), Nouméa, New Caledonia

 Ph.D. student under the co-supervision of Dr. Leigh Torres

Species Distribution Models (SDM), also referred to as ecological niche models, may be defined as “a model that relates species distribution data (occurrence or abundance at known locations) with information on the environmental and/or spatial characteristics of those locations” (Elith & Leathwick, 2009)⁠. In the last couple decades, SDMs have become an indispensable part of the ecologists’ and conservationists’ toolbox. What scientist has not dreamed of being able to summarize a species’ environmental requirements and predict where and when it will occur, all in one tiny statistical model? It sounds like magic… but the short acronym “SDM” is the pretty front window of an intricate and gigantic research field that may extend way beyond the skills of a typical ecologist (even so for a graduate student like myself).

As part of my PhD thesis about the spatial ecology of humpback whales in New Caledonia, South Pacific, I was planning on producing a model to predict their distribution in the region and help spatial planning within the Natural Park of the Coral Sea. An innocent and seemingly perfectly feasible plan for a second year PhD student. To conduct this task, I had at my disposal more than 1,000 sightings recorded during dedicated surveys at sea conducted over 14 years. These numbers seem quite sufficient, considering the rarity of cetaceans and the technical challenges of studying them at sea. And there was more! The NGO Opération Cétacés  also recorded over 600 sightings reported by the general public in the same time period and deployed more than 40 satellite tracking tags to follow individual whale movements. In a field where it is so hard to acquire data, it felt like I had to use it all, though I was not sure how to combine all these types of data, with their respective biases, scales and assumptions.

One important thing about SDM to remember: it is like a cracker section in a US grocery shop, there is sooooo much choice! As I reviewed the possibilities and tested various modeling approaches on my data I realized that this study might be a good opportunity to contribute to the SDM field, by conducting a comparison of various algorithms using cetacean occurrence data from multiple sources. The results of this work was just published  in Diversity and Distributions:

Derville S, Torres LG, Iovan C, Garrigue C. (2018) Finding the right fit: Comparative cetacean distribution models using multiple data sources and statistical approaches. Divers Distrib. 2018;00:1–17. https://doi. org/10.1111/ddi.12782

There are simply too many! Anonymous grocery shops, Corvallis, OR
Credit: Dawn Barlow

If you are a new-comer to the SDM world, and specifically its application to the marine environment, I hope you find this interesting. If you are a seasoned SDM user, I would be very grateful to read your thoughts in the comment section! Feel free to disagree!

So what is the take-home message from this work?

  • There is no such thing as a “best model”; it all depends on what you want your model to be good at (the descriptive vs predictive dichotomy), and what criteria you use to define the quality of your models.

The predictive vs descriptive goal of the model: This is a tricky choice to make, yet it should be clearly identified upfront. Most times, I feel like we want our models to be decently good at both tasks… It is a risky approach to blindly follow the predictions of a complex model without questioning the meaning of the ecological relationships it fitted. On the other hand, conservation applications of models often require the production of predicted maps of species’ probability of presence or habitat suitability.

The criteria for model selection: How could we imagine that the complexity of animal behavior could be summarized in a single metric, such as the famous Akaike Information criterion (AIC) or the Area under the ROC Curve (AUC)? My study, and that of others (e.g. Elith & Graham  H., 2009),⁠ emphasize the importance of looking at multiple aspects of model outputs: raw performance through various evaluation metrics (e.g. see AUCdiff; (Warren & Seifert, 2010)⁠, contribution of the variables to the model, shape of the fitted relationships through Partial Dependence Plots (PDP, Friedman, 2001),⁠ and maps of predicted habitat suitability and associated error. Spread all these lines of evidence in front of you, summarize all the metrics, add a touch of critical ecological thinking to decide on the best approach for your modeling question, and Abracadabra! You end up a bit lost in a pile of folders… But at least you assessed the quality of your work from every angle!

  • Cetacean SDMs often serve a conservation goal. Hence, their capacity to predict to areas / times that were not recorded in the data (which is often scarce) is paramount. This extrapolation performance may be restricted when the model relationships are overfitted, which is when you made your model fit the data so closely that you are unknowingly modeling noise rather than a real trend. Using cross-validation is a good method to prevent overfitting from happening (for a thorough review: Roberts et al., 2017)⁠. Also, my study underlines that certain algorithms inherently have a tendency to overfit. We found that Generalized Additive Models and MAXENT provided a valuable complexity trade-off to promote the best predictive performance, while minimizing overfitting. In the case of GAMs, I would like to point out the excellent documentation that exist on their use (Wood, 2017)⁠, and specifically their application to cetacean spatial ecology (Mannocci, Roberts, Miller, & Halpin, 2017; Miller, Burt, Rexstad, & Thomas, 2013; Redfern et al., 2017).⁠
  • Citizen science is a promising tool to describe cetacean habitat. Indeed, we found that models of habitat suitability based on citizen science largely converged with those based on our research surveys. The main issue encountered when modeling this type of data is the absence of “effort”. Basically, we know where people observed whales, but we do not know where they haven’t… or at least not with the accuracy obtained from research survey data. However, with some information about our citizen scientists and a little deduction, there is actually a lot you can infer about opportunistic data. For instance, in New Caledonia most of the sightings were reported by professional whale-watching operators or by the general public during fishing/diving/boating day trips. Hence, citizen scientists rarely stray far from harbors and spend most of their time in the sheltered waters of the New Caledonian lagoon. This reasoning provides the sort of information that we integrated in our modeling approach to account for spatial sampling bias of citizen science data and improve the model’s predictive performance.

Many more technical aspects of SDM are brushed over in this paper (for detailed and annotated R codes of the modeling approaches, see supplementary information of our paper). There are a few that are not central to the paper, but that I think are worth sharing:

  • Collinearity of predictors: Have you ever found that the significance of your predictors completely changed every time you removed a variable? I have progressively come to discover how unstable a model can be because of predictor collinearity (and the uneasy feeling that comes with it …). My new motto is to ALWAYS check cross-correlation between my predictors, and do it THOROUGHLY. A few aspects that may make a big difference in the estimation of collinearity patterns are to: (1) calculate Pearson vs Spearman coefficients, (2) check correlations between the values recorded at the presence points vs over the whole study area, and (3) assess the correlations between raw environmental variables vs between transformed variables (log-transformed, etc). Though selecting variables with Pearson coefficients < 0.7 is usually a good rule (Dormann et al., 2013), I would worry of anything above 0.5, or at least keep it in mind during model interpretation.
  • Cross-validation: If removing 10% of my dataset greatly impacts the model results, I feel like cross-validation is critical. The concept is based on a simple assumption, if I had sampled a given population/phenomenon/system slightly differently, would I have come to the same conclusion? Cross-validation comes in many different methods, but the basic concept is to run the same model several times (number of times may depend on the size of your data set, hierarchical structure of your data, computation power of your computer, etc.) over different chunks of your data. Model performance metrics (e.g., AUC) and outputs (e.g., partial dependence plots) are than summarized on the many runs, using mean/median and standard deviation/quantiles. It is up to you how to pick these chunks, but before doing this at random I highly recommend reading Roberts et al. (2017).

The evil of the R2: I am probably not the first student to feel like what I have learned in my statistical classes at school is in practice, at best, not very useful, and at worst, dangerously misleading. Of course, I do understand that we must start somewhere, and that learning the basics of inferential statistics is a necessary step to, one day, be able to answer your one research questions. Yet, I feel like I have been carrying the “weight of the R2” for far too long before actually realizing that this metric of model performance (R2 among others) is simply not  enough to trust my results. You might think that your model is robust because among the 1000 alternative models you tested, it is the one with the “best” performance (deviance explained, AIC, you name it), but the model with the best R2 will not always be the most ecologically meaningful one, or the most practical for spatial management perspectives. Overfitting is like a sword of Damocles hanging over you every time you create a statistical model All together, I sometimes trust my supervisor’s expertise and my own judgment more than an R2.

Source: internet

A few good websites/presentations that have helped me through my SDM journey:

General website about spatial analysis (including SDM): http://rspatial.org/index.html

Cool presentation by Adam Smith about SDM:

http://www.earthskysea.org/!ecology/sdmShortCourseKState2012/sdmShortCourse_kState.pdf

Handling spatial data in R: http://www.maths.lancs.ac.uk/~rowlings/Teaching/UseR2012/introductionTalk.html

“The magical world of mgcv”, a great presentation by Noam Ross: https://www.youtube.com/watch?v=q4_t8jXcQgc

 

Literature cited

Dormann, C. F., Elith, J., Bacher, S., Buchmann, C., Carl, G., Carré, G., … Lautenbach, S. (2013). Collinearity: A review of methods to deal with it and a simulation study evaluating their performance. Ecography, 36(1), 027–046. https://doi.org/10.1111/j.1600-0587.2012.07348.x

Elith, J., & Graham  H., C. (2009). Do they? How do they? WHY do they differ? On finding reasons for differing performances of species distribution models . Ecography, 32(Table 1), 66–77. https://doi.org/10.1111/j.1600-0587.2008.05505.x

Elith, J., & Leathwick, J. R. (2009). Species Distribution Models: Ecological Explanation and Prediction Across Space and Time. Annual Review of Ecology, Evolution, and Systematics, 40(1), 677–697. https://doi.org/10.1146/annurev.ecolsys.110308.120159

Friedman, J. H. (2001). Greedy Function Approximation: A gradient boosting machine. The Annals of Statistics, 29(5), 1189–1232. Retrieved from http://www.jstor.org/stable/2699986

Mannocci, L., Roberts, J. J., Miller, D. L., & Halpin, P. N. (2017). Extrapolating cetacean densities to quantitatively assess human impacts on populations in the high seas. Conservation Biology, 31(3), 601–614. https://doi.org/10.1111/cobi.12856.This

Miller, D. L., Burt, M. L., Rexstad, E. A., & Thomas, L. (2013). Spatial models for distance sampling data: Recent developments and future directions. Methods in Ecology and Evolution, 4(11), 1001–1010. https://doi.org/10.1111/2041-210X.12105

Redfern, J. V., Moore, T. J., Fiedler, P. C., de Vos, A., Brownell, R. L., Forney, K. A., … Ballance, L. T. (2017). Predicting cetacean distributions in data-poor marine ecosystems. Diversity and Distributions, 23(4), 394–408. https://doi.org/10.1111/ddi.12537

Roberts, D. R., Bahn, V., Ciuti, S., Boyce, M. S., Elith, J., Guillera-Arroita, G., … Dormann, C. F. (2017). Cross-validation strategies for data with temporal, spatial, hierarchical or phylogenetic structure. Ecography, 0, 1–17. https://doi.org/10.1111/ecog.02881

Warren, D. L., & Seifert, S. N. (2010). Ecological niche modeling in Maxent: the importance of model complexity and the performance of model selection criteria. Ecological Applications, 21(2), 335–342. https://doi.org/10.1890/10-1171.1

Wood, S. N. (2017). Generalized additive models: an introduction with R (second edi). CRC press.