Tag: spatial ecology

Who, where, when: Estimating individual space use patterns of PCFG gray whales

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

Understanding how baleen whales are affected by human activity is a central goal for many research projects in the GEMM Lab. The overarching goal of the GRANITE (Gray whale Response to Ambient Noise Informed by Technology and Ecology) project is to quantify baleen whale physiological response to different stressors (e.g., boat presence and noise) and model the subsequent impacts of these stressors on the population. We will achieve this goal by implementing our long-term, replicate dataset of Pacific Coast Feeding Group (PCFG) gray whales into a framework called population consequences of disturbance (PCoD). I will not go into the details of PCoD in this blog (but I wrote a post a few years ago that you can revisit). Instead, I will explain the approach I am taking to assess where and when individual whales spend time in our study area, which will form an essential component of PCoD and be one of the chapters of my PhD dissertation.

Individuals in a population are unlikely to be exposed to a stressor in a uniform way because they make decisions differently based on intrinsic (e.g., sex, age, reproductive status) and extrinsic (e.g., environment, prey, predators) factors (Erlinge & Sandell 1986). For example, a foraging female gray whale who is still nursing a calf will need to consider factors that are different to ones that an adult single male might need to consider when choosing a location to feed. These differences in decision-making exist across the whole population, which makes it important to understand where individuals are spending time and how they overlap with stressors in space and time before trying to quantify the impacts of stressors on the population as a whole (Pirotta et al. 2018). I am currently working on an analysis that will determine an individual’s exposure to a number of stressors based on their space use patterns. 

We can monitor space use patterns of individuals in a population through time using spatial capture-recapture techniques. As the name implies, a spatial capture-recapture technique involves capturing an individual in a marked location during a sampling period, releasing it back into the population, and then (hopefully) re-capturing it during another sampling period in the future, at either the same or a different location. With enough repeat sampling events, the method should build spatial capture histories of individuals through time to better understand an individual’s space use patterns (Borchers & Efford 2008). While the use of the word capture implies that the animal is being physically caught, this is not necessarily the case. Individuals can be “captured” in a number of non-invasive ways, including by being photographed, which is how we “capture” individual PCFG gray whales. These capture-recapture methods were first pioneered in terrestrial systems, where camera traps (i.e., cameras that take photos or videos when a motion sensor is triggered) are set up in a systematic grid across a study area (Figure 1; Royle et al. 2009, Gray 2018). Placing the cameras in a grid system ensures that there is an equal distribution of cameras throughout the study area, which means that an animal theoretically has a uniform chance of being captured. However, because we know that individuals within a population make space use decisions differently, we assume that individuals will distribute themselves differently across a landscape, which will manifest as individuals having different centers of their spatial activity. The probability of capturing an individual is highest when a camera trap is at that individual’s activity center, and the cameras furthest away from the individual’s activity center will have the lowest probability of capturing that individual (Efford 2004). By using this principle of probability, the data generated from spatial capture-recapture field methods can be modelled to estimate the activity centers and ranges for all individuals in a population. The overlap of an individual’s activity center and range can then be compared to the spatiotemporal distribution of stressors that an individual may be exposed to, allowing us to determine whether and how an individual has been exposed to each stressor. 

Figure 1. Example of camera trap grid in a study area. Figure taken from Gray (2018).

While capture-recapture methods were first developed in terrestrial systems, they have been adapted for application to marine populations, which is what I am doing for our GRANITE dataset of PCFG gray whales. Together with a team of committee members and GRANITE collaborators, I am developing a Bayesian spatial capture-recapture model to estimate individual space use patterns. In order to mimic the camera trap grid system, we have divided our central Oregon coast study area into latitudinal bins that are approximately 1 km long. Unfortunately, we do not have motion sensor activated cameras that automatically take photographs of gray whales in each of these latitudinal bins. Instead, we have eight years of boat-based survey effort with whale encounters where we collect photographs of many individual whales. However, as you now know, being able to calculate the probability of detection is important for estimating an individual’s activity center and range. Therefore, we calculated our spatial survey effort per latitudinal bin in each study year to account for our probability of detecting whales (i.e., the area of ocean in km2 that we surveyed). Next, we tallied up the number of times we observed every individual PCFG whale in each of those latitudinal bins per year, thus creating individual spatial capture histories for the population. Finally, using just those two data sets (the individual whale capture histories and our survey effort), we can build models to test a number of different hypotheses about individual gray whale space use patterns. There are many hypotheses that I want to test (and therefore many models that I need to run), with increasing complexity, but I will explain one here.

Over eight years of field work for the GRANITE project, consisting of over 40,000 km2 of ocean surveyed with 2,169 sightings of gray whales, our observations lead us to hypothesize that there are two broad space use strategies that whales use to optimize how they find enough prey to meet their energetic needs. For the moment, we are calling these strategies ‘home-body’ and ‘roamer’. As the name implies, a home-body is an individual that stays in a relatively small area and searches for food in this area consistently through time. A roamer, on the other hand, is an individual that travels and searches over a greater spatial area to find good pockets of food and does not generally tend to stay in just one place. In other words, we except a home-body to have a consistent activity center through time and a small activity range, while a roamer will have a much larger activity range and its activity center may vary more throughout the years (Figure 2). 

Figure 2. Schematic representing one of the hypotheses we will be testing with our Bayesian spatial capture-recapture models. The schematic shows the activity centers (the circles) and activity ranges (vertical lines attached to the circles) of two individuals (green and orange) across three years in our central Oregon study area. The green individual represents our hypothesized idea of a home-body, whereas the orange individuals represents our idea of a roamer.

While this hypothesis sounds straightforward, there are a lot of decisions that I need to make in the Bayesian modeling process that can ultimately impact the results. For example, do all home-bodies in a population have the same size activity range or can the size vary between different home-bodies? If it can vary, by how much can it vary? These same questions apply for the roamers too. I have a long list of questions just like these, which means a lot of decision-making on my part, and that long list of hypotheses I previously mentioned. Luckily, I have a fantastic team made up of Leigh, committee members, and GRANITE collaborators that are guiding me through this process. In just a few more months, I hope to reveal how PCFG individuals distribute themselves in space and time throughout our central Oregon study area, and hence describe their exposure to different stressors. Stay tuned! 

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References

Borchers DL, Efford MG (2008) Spatially explicit maximum likelihood methods for capture-recapture studies. Biometrics 64:377-385.

Efford M (2004) Density estimation in live-trapping studies. Oikos 106:598-610.

Erlinge S, Sandell M (1986) Seasonal changes in the social organization of male stoats, Mustela erminea: An effect of shifts between two decisive resources. Oikos 47:57-62.

Gray TNE (2018) Monitoring tropical forest ungulates using camera-trap data. Journal of Zoology 305:173-179.

Pirotta E, Booth CG, Costa DP, Fleishman E, Kraus SD, and others (2018) Understanding the population consequences of disturbance. Ecology and Evolution 8(19):9934–9946.Royle J, Nichols J, Karanth KU, Gopalaswamy AM (2009) A hierarchical model for estimating density in camera-trap studies. Journal of Applied Ecology 46:118-127.

How big, how blue, how beautiful! Studying the impacts of climate change on big, (and beautiful) blue whales

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

The SAPPHIRE Project is in full swing, as we spend our days aboard the R/V Star Keys searching for krill and blue whales (Figure 1) in the South Taranaki Bight (STB) region of Aotearoa New Zealand. We are investigating how changing ocean conditions impact krill availability and quality, and how this in turn impacts blue whale behavior, health, and reproduction. Understanding the link between changing environmental conditions on prey species and predators is key to understanding the larger implications of climate change on ocean food webs and each populations’ resiliency. 

Figure 1. The SAPPHIRE team searching for blue whales. Top left) KC Bierlich, top right) Dawn Barlow, bottom left) Dawn Barlow, Kim Bernard (left to right), bottom right) KC Bierlich, Dawn Barlow, Leigh Torres, Mike Ogle (left to right).  

One of the many components of the SAPPHIRE Project is to understand how foraging success of blue whales is influenced by environmental variation (see this recent blog written by Dr. Dawn Barlow introducing each component of the project). When you cannot go to a grocery store or restaurant any time you are hungry, you must rely on stored energy from previous feeds to fuel energy needs. Body condition reflects an individual’s stored energy in the body as a result of feeding and thus represents the foraging success of an individual, which can then affect its potential for reproductive output and the individual’s overall health (see this previous blog). As discussed in a previous blog, drones serve as a valuable tool for obtaining morphological measurements of whales to estimate their body condition. We are using drones to collect aerial imagery of pygmy blue whales to obtain body condition measurements late in the foraging season between years 2024 and 2026 of the SAPPHIRE Project (Figure 2). We are quantifying body condition as Body Area Index (BAI), which is a relative measure standardized by the total length of the whale and well suited for comparing individuals and populations (Figure 3). 

The GEMM Lab recently published an article led by Dr. Dawn Barlow where we investigated the differences in BAI between three blue whale populations: Eastern North Pacific blue whales feeding in Monterey Bay, California; Chilean blue whales feeding in the Corcovado Gulf; and New Zealand Pygmy blue whales feeding in the STB (Barlow et al., 2023). These three populations are interesting to compare since blue whales that feed in Monterey Bay and Corcovado Gulf migrate to and from these seasonally productive feeding grounds, while the Pygmy blue whales stay in Aotearoa New Zealand year-round. Interestingly, the Pygmy blue whales had higher BAI (were fatter) compared to the other two regions despite relatively lower productivity in their foraging grounds. This difference in body condition may be due to different life history strategies where the non-migratory Pygmy blue whales may be able to feed as opportunities arrive, while the migratory strategies of the Eastern North Pacific and Chilean blue whales require good timing to access high abundant prey. Another interesting and unexpected result from our blue whale comparison was that Pygmy blue whales are not so “pygmy”; they are actually the same size as Eastern North Pacific and Chilean blue whales, with an average size around 22 m. Our findings from this blue whale comparison leads us to more questions about how environmental conditions that vary from year to year influence body condition and reproduction of these “not so pygmy” blue whales. 

Figure 2. An aerial image of a Pygmy blue whale in the South Taranaki Bight region of Aotearoa New Zealand collected during the SAPPHIRE 2024 field season using a DJI Inspire 2 drone. 
Figure 3. A drone image of a Pygmy blue whale and the length and body width measurements used to estimate Body Area Index (BAI), represented by the shaded blue region. Width measurements will also be used to help identify pregnant individuals.

The GEMM Lab has been studying this population of Pygmy blue whales in the STB since 2013 and found that years designated as a marine heatwave resulted with a reduction in blue whale feeding activity. Interestingly, breeding activity is also reduced during marine heatwaves in the following season when compared to the breeding season following a more productive, typical foraging season. These findings indicate that fluctuations in the environment, such as marine heatwaves, may affect not only foraging success, but also reproduction in Pygmy blue whales. 

To help us better understand reproductive patterns across years, we will use body width measurements from drone images paired with hormone concentrations collected from fecal and biopsy samples to identify pregnant individuals. Progesterone is a hormone secreted in the ovaries of mammals during the estrous cycle and gestation, making it the predominant hormone responsible for sustaining pregnancy. Recently, the GEMM Lab’s Dr. Alejandro Fernandez-Ajo wrote a blog discussing his publication identifying pregnant individual gray whales using drone-based body width measurements and progesterone concentrations from fecal samples (Fernandez et al., 2023). While individuals that were pregnant had higher levels of progesterone compared to when they were not pregnant, the body width at 50% of the body length served as a more reliable method for detecting pregnancy in gray whales. We will use similar methods to help identify pregnancy in Pygmy blue whales for the SAPPHIRE Project where will we examine body width measurement paired with progesterone concentrations collected from fecal and biopsy samples to identify pregnant individuals. We hope our work will help to better understand how climate change will influence Pygmy blue whale body condition and reproduction, and thus the overall health and resiliency of the population. Stay tuned! 

References

Barlow, D. R., Bierlich, K. C., Oestreich, W. K., Chiang, G., Durban, J. W., Goldbogen, J. A., Johnston, D. W., Leslie, M. S., Moore, M. J., Ryan, J. P., & Torres, L. G. (2023). Shaped by Their Environment: Variation in Blue Whale Morphology across Three Productive Coastal Ecosystems. Integrative Organismal Biology, 5(1). https://doi.org/10.1093/iob/obad039

Fernandez Ajó, A., Pirotta, E., Bierlich, K. C., Hildebrand, L., Bird, C. N., Hunt, K. E., Buck, C. L., New, L., Dillon, D., & Torres, L. G. (2023). Assessment of a non-invasive approach to pregnancy diagnosis in gray whales through drone-based photogrammetry and faecal hormone analysis. Royal Society Open Science, 10(7), 230452. https://doi.org/10.1098/rsos.230452

Making predictions: A window into ecological forecast models

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

“What is the weather going to be like tomorrow?” “How long will it take to drive there, with traffic?” We all rely on forecasts to make decisions, such as whether to bring a rain jacket, when to get in the car to arrive at a certain destination on time, or any number of situations where we want a prediction of what will happen in the near future. Statistical models underpin many of these examples, using past data to inform future predictions.

Early on in graduate school, I was told that “all models are wrong, but some models work.” Any model is essentially a best approximation, using mathematical relationships, of how we understand a pattern. Models are powerful tools in ecology, enabling us to distill complex, dynamic, and interacting systems into terms and parameters that can be quantified. This ability can help us better understand our study systems and use that understanding to make predictions. We will never be able to describe every nuance of an ecosystem. Instead, the challenge is to collect enough information to build an informed model that can enhance our understanding, without over-simplifying or unnecessarily complicating the system we aim to describe. As Dr. Simon Levin stated in his 1989 seminal paper:

“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.”1

Species distribution models (SDMs) are the particular branch of models that underpin much of my PhD research on blue whale ecology and distribution in New Zealand. 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)2. The model is a best attempt to quantify and describe the relationships between predictors, e.g., temperature and the observed species distribution pattern. For example, blue whale occurrence is higher in areas of lower temperatures and greater krill availability, and these relationships can be described with models3. So, a model essentially takes all the data available, and synthesizes that information in terms of the relationships between the predictors (environment) and response (species occurrence). Then, we can look at the fitted relationships to ask what we would expect from the species distribution pattern when temperature, or krill availability, or any other predictor, is at a particular value. 

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).

So, if a model is simply a mathematical description of how terms interact to produce a particular outcome, how do predictions work? To make a spatial prediction, e.g., a map of the probability of a species being present, you need two things: a model describing the functional relationships between species presence and your environmental predictors, and the values of your predictor variables on the day you are interested in predicting to. For example, you may need to obtain a map of sea surface temperature, productivity, temperature anomaly, and surface currents on a day you want to know where whales are expected to be. Your model is the applied across that stack of spatial environmental layers and, based on the functional relationships derived by the model, you get an estimate of the probability of species occurrence based on the temperature, productivity, anomaly, and surface current values at each location. By applying the model over a range of values, you can obtain a continuous surface with the probability of presence, in the form of a map. These maps are typically for the past or present because that is when we can typically acquire spatial environmental layers. However, to make predictions for a future time of interest, we need to have spatial environmental layers for the future.

Forecasts are predictions for the future. Recent advances in technology and computing have led to an emergence of environmental and ecological forecasting tools that are being developed around the world to produce marine forecasts. These tools include predictions of the physical environment such as ocean temperatures or currents, and biological patterns such as where species will be distributed in space and timing of events like salmon spawning or lobster landings4. The ability to generate forecast of marine ecosystems is of particular interest to resource users and managers because it can allow them to be proactive rather than reactive. Forecasts enable us to anticipate events or patterns and prepare, rather than having to respond in real-time or after the fact.

The South Taranaki Bight region in New Zealand is an area where blue whale foraging habitat frequently coincides with industry pressures, including petroleum and mineral extraction, exploration for petroleum reserves using seismic airgun surveys, vessel traffic between ports, and even an ongoing proposal for seabed mining5. Static spatial restrictions to mitigate impacts from these activities on blue whales may be met with resistance from industry user groups, but dynamic spatial management6–8 of blue whale habitat could be more attractive and acceptable. The key for successful dynamic management is knowing where and when to put those boundaries; and this is where ecological forecast models can show their strength. If we can predict suitable blue whale habitat for the future, proactive regulations can be applied to enhance conservation management in the region. Can we develop reliable and useful ecological forecasts for the South Taranaki Bight? Well, given that we have already developed robust models of the relationships between blue whales and their habitat3 and have documented the spatial and temporal lags between wind, upwelling, and blue whales9, we feel confident that we can develop forecast models to predict where blue whales will be in the STB region. As we continue working hard toward this goal, we invite you to check back for our findings in the future. So, consider this blog post a forecast of sorts, and stay tuned!  

Figure 2. A blue whale surfaces in front of an oil extraction platform in the South Taranaki Bight, demonstrating the overlap between whales and industry in the region. Photo by D. Elvines.

References:

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

2.        Elith, J. & Leathwick, J. R. Species Distribution Models: Ecological Explanation and Prediction Across Space and Time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697 (2009).

3.        Barlow, D. R., Bernard, K. S., Escobar-Flores, P., Palacios, D. M. & Torres, L. G. 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. 642, 207–225 (2020).

4.        Payne, M. R. et al. Lessons from the first generation of marine ecological forecast products. Front. Mar. Sci. 4, 1–15 (2017).

5.        Torres, L. G. Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zeal. J. Mar. Freshw. Res. 47, 235–248 (2013).

6.        Hyrenbach, K. D., Forney, K. A. & Dayton, P. K. Marine protected areas and ocean basin management. Aquat. Conserv. Mar. Freshw. Ecosyst. 10, 437–458 (2000).

7.        Maxwell, S. M. et al. Dynamic ocean management: Defining and conceptualizing real-time management of the ocean. Mar. Policy 58, 42–50 (2015).

8.        Oestreich, W. K., Chapman, M. S. & Crowder, L. B. A comparative analysis of dynamic management in marine and terrestrial systems. Front. Ecol. Environ. 18, 496–504 (2020).

9.        Barlow, D. R., Klinck, H., Ponirakis, D., Garvey, C. & Torres, L. G. Temporal and spatial lags between wind, coastal upwelling, and blue whale occurrence. Sci. Rep. 11, (2021).