The right tool for the job: examining the links between animal behavior, morphology and habitat

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

In order to understand a species’ distribution, spatial ecologists assess which habitat characteristics are most often associated with a species’ presence. Incorporating behavior data can improve this analysis by revealing the functional use of each habitat type, which can help scientists and managers assign relative value to different habitat types. For example, habitat used for foraging is often more important than habitat that a species just travels through. Further complexity is added when we consider that some species, such as gray whales, employ a variety of foraging tactics on a variety of prey types that are associated with different habitats. If individual foraging tactic specialization is present, different foraging habitats could be valuable to specific subgroups that use each tactic. Consequently, for a population that uses a variety of foraging tactics, it’s important to study the associations between tactics and habitat characteristics.

Lukoschek and McCormick’s (2001) study investigating the spatial distribution of a benthic fish species’ foraging behavior is a great example of combining data on behavior, habitat, and morphology.  They collected data on the diet composition of individual fish categorized into different size classes (small, medium, and large) and what foraging tactics were used in which reef zones and habitat types. The foraging tactics ranged from feeding in the water column to digging (at a range of depths) in the benthic substrate. The results showed that an interesting combination of fish behavior and morphology explained the observed diet composition and spatial distribution patterns. Small fish foraged in shallower water, on smaller prey, and primarily employed the water column and shallow digging tactics. In contrast, large fish foraged in deep water, on larger prey, and primarily fed by digging deeper into the seafloor (Figure 1). This pattern is explained by both morphology and behavior. Morphologically, the size of the feeding apparatus (mouth gape size) affects the size of the prey that a fish can feed on. The gape of the small fish is not large enough to eat the larger prey that large fish are able to consume. Behaviorally, predation risk also affects habitat selection and tactic use. Small fish are at higher risk of being predated on, so they remain in shallow areas where they are more protected from predators and they don’t dig as deep to forage because they need to be able to keep an eye out for predators. Interestingly, while they found a relationship between the morphology of the fish and habitat use, they did not find an association between specific feeding tactics and habitat types.

Figure 1. Figure from Lukoschek and McCormick (2001) showing that small fish (black bar) were found in shallow habitat while large fish (white bar) were found in deep habitat.

Conversely, Torres and Read (2009) did find associations between theforaging tactics of bottlenose dolphins in Florida Bay, FL and habitat type. Dolphins in this bay employ three foraging tactics: herd and chase, mud ring feeding, and deep diving. Observations of the foraging tactics were linked to habitat characteristics and individual dolphins. The study found that these tactics are spatially structured by depth (Figure 2), with deep diving occurring in deep water whereas mud ring feeding occurrs in shallower water. They also found evidence of individual specialization! Individuals that were observed deep diving were not observed mud ring feeding and vice-versa. Furthermore, they found that individuals were found in the habitat type associated with their preferred tactic regardless of whether they were foraging or not. This result indicates that individual dolphins in this bay have a foraging tactic they prefer and tend to stay in the corresponding habitat type. These findings are really intriguing and raise interesting questions regarding how these tactics and specializations are developed or learned. These are questions that I am also interested in asking as part of my thesis.

Figure 2. Figure from Torres and Read (2009) showing that deep diving is associated with deeper habitat while mud ring feeding is associated with shallow habitat.

Both of these studies are cool examples that, combined, exemplify questions I am interested in examining using our study population of Pacific Coast Feeding Group (PCFG) gray whales. Like both studies, I am interested in assessing how specific foraging tactics are associated with habitat types. Our hypothesis is that different prey types live in different habitat types, so each tactic corresponds to the best way to feed on that prey type in that habitat. While predation risk doesn’t have as much of an effect on foraging gray whales as it does on small benthic fish, I do wonder how disturbance from boats could similarly affect tactic preference and spatial distribution. I am also curious to see if depth has an effect on tactic choice by using the morphology data from our drone-based photogrammetry. Given that these whales forage in water that is sometimes as deep as they are long, it stands to reason that maneuverability would affect tactic use. As described in a previous blog, I’m also looking for evidence of individual specialization. It will be fascinating to see how foraging preference relates to space use, habitat preference, and morphology.

These studies demonstrate the complexity involved in studying a population’s relationship to its habitat. Such research involves considering the morphology and physiology of the animals, their social, individual, foraging, and predator-prey behaviors, and the relationship between their prey and the habitat. It’s a bit daunting but mostly really exciting because better understanding each puzzle piece improves our ability to estimate how these animals will react to changing environmental conditions.

While I don’t have any answers to these questions yet, I will be working with a National Science Foundation Research Experience for Undergraduates intern this summer to develop a habitat map of our study area that will be used in this analysis and potentially answer some preliminary questions about PCFG gray whale habitat use patterns. So, stay tuned to hear more about our work this summer!


Lukoschek, V., & McCormick, M. (2001). Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): Relationship between foraging behaviour, habitat use, jaw size, and prey selection. Marine Biology, 138(6), 1099–1113.

Torres, L. G., & Read, A. J. (2009). Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins ( Tursiops truncatus ) in Florida Bay, Florida. Marine Mammal Science, 25(4), 797–815.

Dolphin Diets: Common bottlenose dolphin prey preferences off California

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

Humans are fascinated by food. We want to know its source, its nutrient content, when it was harvested and by whom, and so much more. Since childhood, I was the nagging child who interrogated wait staff about the seafood menu because I cared about the sustainability aspect as well as consuming ethically-sourced seafood. Decades later I still do the same: ask a myriad of questions from restaurants and stores in order to eat as sustainably as possible. But in addition to asking these questions about my food, I also question what my study species eats and why. My study populations, common bottlenose dolphins, are described as top opportunistic predators (Norris and Prescott 1961, Shane et al. 1986, Barros and Odell 1990). In my study area off of California, this species exists in two ecotypes. The coastal ecotype off of California, USA are generalist predators, feeding on many different species of fish using different foraging techniques (Ballance 1992, Shane 1990). The offshore ecotype, on the other hand, is less well-studied, but is frequently observed in association with sperm whales, although the reason is still unknown (Díaz-Gamboa et al. 2018). Stable isotope analysis from skin samples from the two ecotypes indicates that the ecotypes exhibit different foraging strategies based on different isotopic carbon and nitrogen levels (Díaz-Gamboa et al. 2018).

Growing up, I kept the Monterey Bay Aquarium’s Seafood Watch Guide with me to choose the most sustainably-sourced seafood at restaurants. Today there is an easy-to-use application for mobile phones that replaced the paper guide. (Image Source:

Preliminary and historical data on common bottlenose dolphins (Tursiops truncatus) suggest that the coastal ecotype spend more time near estuary mouths than offshore dolphins (Ballance 1992, Kownacki et al. unpublished data). Estuaries contain large concentrations of nutrients from runoff, which support zooplankton and fishes. It is for this reason that these estuaries are thought to be hotspots for bottlenose dolphin foraging. Some scientists hypothesize that these dolphins are estuarine-based prey specialists (Barros and Odell 1990), or that the dolphins simply aggregate in estuaries due to higher prey abundance (Ballance 1992).

Coastal bottlenose dolphins traveling near an estuary mouth in San Diego, CA. (Photographed under NOAA NMFS Permit # 19091).

In an effort to understand diet compositions of bottlenose dolphins, during coastal surveys seabirds were recorded in association with feeding groups of dolphins. Therefore, it is reasonable to believe that dolphins were feeding on the same fishes as Brown pelicans, blue-footed and brown boobies, double-crested cormorants, and magnificent frigatebirds, seeing as they were the most common species associated with bottlenose dolphin feeding groups (Ballance 1992). A shore-based study by Hanson and Defran (1993) found that coastal dolphins fed more often in the early morning and late afternoon, as well as during periods of high tide current. These patterns may have to do with the temporal and spatial distribution of prey fish species. From the few diet studies conducted on these bottlenose dolphins in this area, 75% of the prey were species from the families Ebiotocidae (surf perches) and Sciaendae (croakers) (Norris and Prescott 1961, Walker 1981). These studies, in addition to optimal foraging models, suggest this coastal ecotype may not be as much of a generalist as originally suggested (Defran et al. 1999).

A redtail surfperch caught by a fisherman from a beach in San Diego, CA. These fish are thought to be common prey of coastal bottlenose dolphins. (Image Source: FishwithJD)

Diet studies on the offshore ecotype of bottlenose dolphins worldwide show a preference for cephalopods, similar to other toothed cetaceans who occupy similar regions, such as Risso’s dolphin, sperm whales, and pilot whales (Clarke 1986, Cockcroft and Ross 1990, Gonzalez et al. 1994, Barros et al. 2000, Walker et al. 1999). Because these animals seldom strand on accessible beaches, stomach contents analyses are limited to few studies and isotope analysis is more widely available from biopsies. We know these dolphins are sighted in deeper waters than the habitat of coastal dolphins where there are fewer nutrient plumes, so it is reasonable to hypothesize that the offshore ecotype consumes different species and may be more specialized than the coastal ecotype.

An bottlenose dolphin forages on an octopus. (Image source: Mandurah Cruises)

For a species that is so often observed from shore and boats, and is known for its charisma, it may be surprising that the diets of both the coastal and offshore bottlenose dolphins are still largely unknown. Such is the challenge of studying animals that live and feed underwater. I wish I could simply ask a dolphin, much like I would ask staff at restaurants: what is on the menu today? But, unfortunately, that is not possible. Instead, we must make educated hypotheses about the diets of both ecotypes based on necropsies and stable isotope studies, and behavioral and spatial surveys. And, I will continue to look to new technologies and creative thinking to provide the answers we are seeking.

Literature cited:

Ballance, L. T. (1992). Habitat use patterns and ranges of the bottlenose dolphin in the Gulf of California, Mexico. Marine Mammal Science8(3), 262-274.

Barros, N.B., and D. K. Odell. (1990). Food habits of bottlenose dolphins in the southeastern United States. Pages 309-328 in S. Leatherwood and R. R. Reeves, eds. The bottlenose dolphin. Academic Press, San Diego, CA.

Barros, N., E. Parsons and T. Jefferson. (2000). Prey of bottlenose dolphins from the South China Sea. Aquatic Mammals 26:2–6.

Clarke, M. 1986. Cephalopods in the diet of odontocetes. Pages 281–321 in M. Bryden and R. Harrison, eds. Research on dolphins. Clarendon Press, Oxford, NY.

Cockcroft, V., and G. Ross. (1990). Food and feeding of the Indian Ocean bottlenose dolphin off southern Natal, South Africa. Pages 295–308 in S. Leatherwood and R. R. Reeves, eds. The bottlenose dolphin. Academic Press, San Diego, CA.

Defran, R. H., Weller, D. W., Kelly, D. L., & Espinosa, M. A. (1999). Range characteristics of Pacific coast bottlenose dolphins (Tursiops truncatus) in the Southern California Bight. Marine Mammal Science15(2), 381-393.

Díaz‐Gamboa, R. E., Gendron, D., & Busquets‐Vass, G. (2018). Isotopic niche width differentiation between common bottlenose dolphin ecotypes and sperm whales in the Gulf of California. Marine Mammal Science34(2), 440-457.

Gonzalez, A., A. Lopez, A. Guerra and A. Barreiro. (1994). Diets of marine mammals stranded on the northwestern Spanish Atlantic coast with special reference to Cephalopoda. Fisheries Research 21:179–191.

Hanson, M. T., and Defran, R. H. (1993). The behavior and feeding ecology of the Pacific coast bottlenose dolphin, Tursiops truncatus. Aquatic Mammals19, 127-127.

Norris, K. S., and J. H. Prescott. (1961). Observations on Pacific cetaceans of Californian and Mexican waters. University of California Publications of Zoology 63:29, 1-402.

Shane, S. H. (1990). Comparison of bottlenose dolphin behavior in Texas and Florida, with a critique of methods for studying dolphin behavior. Pages 541-558 in S. Leatherwood and R. R. Reeves, eds. The bottlenose dolphin. Academic Press, San Diego, CA.

Shane, S., R. Wells and B. Wursig. (1986). Ecology, behavior and social organization of bottlenose dolphin: A review. Marine Mammal Science 2:34–63.

Walker, W.A. (1981). Geographical variation in morphology and biology of the bottlenose dolphins (Tursiops) in the eastern North Pacific. NMFS/SWFC Administrative Report. No, LJ-91-03C.

Walker, J., C. Potter and S. Macko. (1999). The diets of modern and historic bottlenose dolphin populations reflected through stable isotopes. Marine Mammal Science 15:335–350.

Dual cameras provide bigger picture

By Hunter Warick, Research Technician, Geospatial Ecology of Marine Megafauna Lab, Marine Mammal Institute

When monitoring the health of a capital breeding species, such as whales that store energy to support reproduction costs, it is important to understand what processes and factors drive the status of their body condition. Information gained will allow for better insight into their cost of reproduction and overall life history strategies.

For the past four years the GEMM Lab has utilized the perspective that Unoccupied Aerial Systems (UAS; or ‘drones’) provide for observations of marine mammals. This aerial perspective has documented gray whale behavior such as jaw snapping, drooling mud, and headstands, all of which shows or suggest foraging (Torres et al. 2018). However, UAS is limited to a bird’s eye view, allowing us to see WHAT whales are doing, but limited information about the reasons WHY. To overcome this hurdle, Leigh Torres and team have equipped their marine mammal research utility belts with the use of GoPro cameras. They developed a technique known as the “GoPro drop” where a GoPro camera mounted to a weighted pole is lowered off the side of the research vessel in waters < 20 m deep via a line to record video data. This technique allows the team to obtain fine-scale habitat and prey variation information, like what the whale experiences. Along with the context provided by the UAS, this dual camera perspective allows for deeper insight into gray whale foraging strategies and efficiency. Torres’s GoPro data analysis protocol examines kelp density, kelp health, benthic substrate, rock fish density, and mysid density. These characteristics are graded along a scale (Figure 1), allowing for relative comparisons of habitat and prey availability between where whales spend time and forage. These GoPro drops will also help create a fine-scale benthic habitat map of the Newport field area. So, why are these data on gray whale habitat and prey important to understand?

Figure 1. The top row shows varying degrees of mysid density (low to high, left to right). Middle row illustrates different types of substrate you might encounter (reef, sandy, boulders; left to right). Bottom row shows the different levels of kelp health (poor, medium, good).

The foraging grounds are the first step in the life history domino chain reaction for many rorqual whales; if this step doesn’t go off cleanly then everything else fails to fall into place. Gray whales partake on a 15,000-20,000 km (round trip) migration, which is the longest of any known mammal (Swartz 1986). During this migration, whales spend around three months fasting in their breeding grounds (Highsmith & Coyle 1992), living only off the energy stores that they accumulated in their feeding grounds (Næss et al. 1998). These extreme conditions of existence for gray whales drive the need to be a successful forager and is why it is so crucial for them to forage in high prey density areas (Newell, C. 2009).

Mysids are a critical part of the gray whale diet in Oregon waters (Newell, C. 2009; Sullivan, F. 2017) and mysids have strong predator-prey relationships with both top-down and bottom-up control (Dunham & Duffus 2001; Newell & Cowles 2006). This unique tie illustrates the great dependency that gray whales have on mysids, further showing the benefit to looking at the density of mysids where gray whales are seen foraging. The quality of mysids may also be as important as quantity; with higher water temperatures resulting in lower lipid content in mysids (Mauchline 1980), suggesting density might not be the only factor for determining efficient whale foraging. The overall goal of gray whales on their foraging grounds is to get as fat as possible in order to reproduce as often as possible. But, this isn’t always as easy as it sounds. Gray whales typically have a two-year breeding interval but can be anywhere from 1-4 years (Blokhin 1984). The longer time it takes to build up adequate energy stores to support reproduction costs, the longer it will take to breed successfully. Building back up these energy stores can prove to be difficult, especially for lactating females (Figure 2).

Being able to track the health and behavior of gray whales on an individual level, including comparisons between variation in body condition, foraging behavior, and fine scale information on benthic communities gained through the use of GoPros, can provide a better understanding of the driving factors and impacts on their health and population trends (Figure 3).

Figure 3. A compilation of video clips captured by the GEMM Lab during their research on gray whale ecology and physiology off Newport, Oregon using Unoccupied Aerial Systems (UAS, or “drones”) and GoPro cameras. UAS are used to observe gray whale behavior and conduct photogrammetry assessment of body condition. GoPro camera drops assess the benthic habitat and prey density across the study region, with a couple chance encounters of whales. Research is conducted under NOAA/NMFS permit # 21678.

The Seascape of Fear: What are the ecological implications of being afraid in the marine environment?

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

In the GEMM Lab, our research focuses largely on the ecology of marine top predators. Inherent in our work are often assumptions that our study species—wide-ranging predators including whales, dolphins, otters, or seabirds—will distribute themselves relative to their prey. In order to make a living in the highly patchy and dynamic marine environment, predators must find ways to predictably locate and exploit prey resources.

But what about the prey? How do the prey structure themselves relative to their predators? This question is explored in depth in a paper titled “The Landscape of Fear: Ecological Implications of Being Afraid” (Laundre et al. 2010), which we discussed in our most recent lab meeting. When wolves were re-introduced in Yellowstone, the elk increased their vigilance and altered their grazing patterns. As a result, the plant community was altered to reflect this “landscape of fear” that the elk move through, where their distribution not only reflected opportunities for the elk to eat but also the risk of being eaten.

Translating the landscape of fear concept to the marine environment is tricky, but a fascinating exercise in ecological theory. We grappled with drawing parallels between the example system of wolves, elk, and vegetation and baleen whales, zooplankton, and phytoplankton. Relative to grazing mammals like elk, the cognitive abilities of zooplankton like krill, copepods, and mysid might pale in comparison. How could we possibly measure “fear” or “vigilance” in zooplankton? The swarming behavior of mysid and krill into dense patches is a defense mechanism—the strategy they have evolved to lessen the likelihood that any one of them will be eaten by a predator. I would posit that the diel vertical migration (DVM) of zooplankton is a manifestation of fear, at least on some level. DVM occurs over the course of each day, with plankton in pelagic ecosystems migrating vertically in the water column to avoid predators by hiding at depth during the daylight hours, and then swimming upward to feed on phytoplankton under the cover of darkness. I won’t speculate any further on the intelligence of zooplankton, but the need to survive predation has driven them to evolve this effective evolutionary strategy of hiding in the ocean’s twilight zone, swimming upward to feed only after dark so that they’re less likely to linger in spaces occupied by predators.

Laundre et al. (2010) present a visual representation of the landscape of fear (Fig. 1, reproduced below), where as an animal moves through space (represented as distance in meters or kilometers, for example), they also move through varying levels of predation risk. Environmental gradients (temperature, for example) tend to be much more stable across space in terrestrial ecosystems such as in the Yellowstone example from the paper. I wonder whether the same concept and visual depiction of a landscape of fear could be translated as risk across various environmental gradients, rather than geographic distances? In this proposed illustration, a landscape of fear would vary based on gradients of environmental conditions rather than geographic space. Such a shift in spatial reference —from geographic to environmental space—might make the model more applicable in the dynamic ocean ecosystems that we study.

What about cases when the predators we study become prey? One example we discussed was gray whales migrating from breeding lagoons in Mexico to feeding grounds in the Bering Sea. Mother-calf pairs hug the coastline tightly, by no means taking the most direct route between locations and adding considerable travel distance to their migration. The leading hypothesis is that mother gray whales take the coastal route to minimize the risk that their calves will fall prey to killer whale attacks. Are there other cases where the predators we study operate in a seascape of fear that we do not yet understand? Likely so, and the predators’ own seascape of fear may account for cases when we cannot explain predator distribution simply by their prey and their environment. To take this a step further, it might be beneficial not only to think of predation risk as only the potential to be eaten, but expand our definition to include human disturbance. While humans may not directly prey on marine predators, the disturbance from human activity in the ocean likely creates a layer of fear which animals must navigate, even in the absence of actual predation.

Our lively lab meeting discussion prompted me to look into how the landscape of fear model has been applied to the highly dynamic and intricate marine environment. In a study examining predator-prey dynamics of three species of marine mammals—bottlenose dolphins, harbor seals, and dugongs—Wirsing et al. (2007) found that in all three cases, the study species spent less time in more desirable prey patches or decreased riskier behavior in the presence of predators. Most studies in marine ecology are observational, as we rarely have the opportunity to manipulate our study system for experimental design and hypothesis testing. However, a study of coral reefs in the Florida Keys conducted by Catano et al. (2015) used fabricated predators—decoys of black grouper, a predatory fish—to investigate the influence of fear of predation on the reef system. What they found was that herbivorous fish consumed significantly less and fed at a much faster rate in the presence of this decoy predator. The grouper, even in decoy form, created a “reefscape of fear”, altering patterns in herbivory with potential ramifications for the entire ecosystem.

My takeaway from our discussion and my musings in this week’s blog post is that predator and prey distribution and behavior is highly interconnected. While predators distribute themselves to maximize their ability to find a meal, their prey respond accordingly by balancing finding a meal of their own with minimizing the risk that they will be eaten. Ecology is the study of an ecosystem, which means the questions we ask are complicated and hierarchical, and must be considered from multiple angles, accounting for biological, environmental, and behavioral elements to name a few. These challenges of studying ecosystems are simultaneously what make ecology fascinating, and exciting.


Laundré, J. W., Hernández, L., & Ripple, W. J. (2010). The landscape of fear: ecological implications of being afraid. Open Ecology Journal3, 1-7.

Catano, L. B., Rojas, M. C., Malossi, R. J., Peters, J. R., Heithaus, M. R., Fourqurean, J. W., & Burkepile, D. E. (2016). Reefscapes of fear: predation risk and reef hetero‐geneity interact to shape herbivore foraging behaviour. Journal of Animal Ecology85(1), 146-156.

Wirsing, A. J., Heithaus, M. R., Frid, A., & Dill, L. M. (2008). Seascapes of fear: evaluating sublethal predator effects experienced and generated by marine mammals. Marine Mammal Science24(1), 1-15.

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.


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.


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.

Understanding sea otter effects through complexity

By Dominique Kone, Masters Student in Marine Resource Management

Species reintroductions are a management strategy to augment the reestablishment or recovery of a locally-extinct or extirpated species into once native habitat. The potential for reestablishment success often depends on the species’ ecological characteristics, habitat requirements, and relationship and effects to other species in the environment[1]. While the science behind species reintroductions is continuously evolving and improving, reintroductions are still inherently risky and uncertain in nature. Therefore, every effort should be made to fully assess ecological factors before a reintroduction takes place. As Oregon considers a potential sea otter reintroduction, understanding these ecological factors is an important piece of my own graduate research.

Sea otters are oftentimes referred to as keystone species because they can have wide-reaching effects on the community structure and function of nearshore marine environments. Furthermore, relative to other marine mammals or top predators, several papers have documented these effects – partially due to the ease in observing their foraging and social behaviors, which typically take place close to shore. In many of these studies, a classic paradigm repeatedly appears: when sea otters are present, prey densities (e.g., sea urchins) are significantly reduced, while macroalgae (e.g., kelp, seagrass) densities are high.

Source: Belleza.

While this paradigm is widely-accepted amongst researchers, a few key studies have also demonstrated that the effects of sea otters may be more variable than we once thought. The paradigm does not necessarily hold true everywhere sea otters exist, or at least not to the same degree. For example, after observing benthic communities along islands with varying sea otter densities in the Aleutian archipelago, Alaska, researchers found that islands with abundant otter populations consistently supported low sea urchin densities and high, yet variable, kelp densities. In contrast, islands without otters consistently had low kelp densities and high, yet variable, urchin densities[2]. This study demonstrates that while the classic paradigm generally held true, the degree to which the ecosystem belonged to one of two dominant states (sea otters, low urchins, and high kelp or no sea otters, high urchins, and low kelp) was less obvious.

This example demonstrates the danger in applying this one-size-fits-all paradigm to sea otter effects. Hence, we want to achieve a better understanding of potential sea otter effects so that managers may anticipate how Oregon’s nearshore environments may be affected if sea otters were to be reintroduced. Yet, how can we accurately anticipate these effects given these potential variations and deviations from the paradigm? Interestingly, if we look to other fields outside ecology, we find a possible solution and tool for tackling these uncertainties: a systematic review of available literature.

Two ecosystem states as predicted by the classic paradigm (left: kelp-dominated; right: urchin-dominated). Source:

For decades, medical researchers have been conducting systematic reviews to assess the efficacy of treatments and drugs by combining several studies to find common findings[3]. These findings can then be used to determine any potential variation between studies (i.e. instances where the results may conflict or differ from one another) and even test the influence and importance of key factors that may be driving that variation[4]. While systematic reviews are quite popular within the medical research field, they have not been applied regularly in ecology, but recognition of their application to ecological questions is growing[5]. In our case of achieving a better understanding of the drivers of ecological impacts of sea otter, a systematic literature review is an ideal tool to assess variable effects. This review will be the focus of my second thesis chapter.

In conducting my review, there will be three distinct phases: (1) review design and study collection, (2) meta-analysis, and (3) factor testing. In the first phase (review design and study collection), I will search the existing literature to collect studies that explicitly compare the availability of key ecosystem components (i.e. prey species, non-prey species, and macroalgae species) when sea otters are absent and present in the environment. By only including studies that make this comparison, I will define effects as the proportional change in each species’ or organism group’s availability (e.g. abundance, biomass, density, etc.) with and without sea otters. In determining these effects, it’s important to recognize that sea otters alter ecosystems via both direct and indirect pathways. Direct effects can be thought of as any change to prey availability via sea otter predation directly, while indirect effects can be thought of an any alteration to the broader ecosystem (i.e. non-prey species, macroalgae, habitat features) as an indirect result from sea otter predation on prey species. I will record both types of effects.

General schematic of a meta-analysis in a systematic review. A meta-analysis is the process of taking multiple datasets (i.e. Data 1, Data 2 etc.) from literature sources, calculating summary statistics or effects (i.e. Summary 1, Summary 2, etc.) for each dataset, running statistical procedures (e.g. SMA = sequential meta-analysis) to relate summary effects and investigate between study variation, and identifying important features driving variation. Source: MediCeption.

In phase two, I will use meta-analytical procedures (i.e. statistical analyses specific to systematic reviews) to calculate one standardized metric to represent sea otter effects. These effects will be calculated and averaged across all collected studies. As previously discussed, there may be key factors – such as sea otter density – that influence these effects. Therefore, in phase three (factor testing), effects will also be calculated separately for each a priori factor to test their influence on the effects. Such factors may include habitat type (i.e. hard or soft sediment), prey species (i.e. sea urchins, crabs, clams, etc.), otter density, depth, or time after otter recolonization.

In statistical terms, the goal of testing factors is to see if the variation between studies is impacted by calculating sea otter effects separately for each factor versus across all studies. In other words, if we find high variation in effects between studies, there may be important factors driving that variation. Therefore, in systematic reviews, we recalculate effects separately for each factor to try to explain that variation. If, however, after testing these factors, variation remains high, there may be other factors that we didn’t test that could be driving that remaining variation. Yet, without a priori knowledge on what those factors could be, such variation should be reported as a major source of uncertainty.

Source: Giancarlo Thomae.

Predicting or anticipating the effects of reintroduced species is no easy feat. In instances where the ecological role of a species is well known – and there is adequate data – researchers can develop and use ecosystem models to predict with some certainty what these effects may be. Yet, in other cases where the species’ role is less studied, has less data, or is more variable, researchers must look to other tools – such as systematic reviews – to gain a better understanding of these potential effects. In this case, a systematic review on sea otter effects may prove particularly useful in helping managers understand what types of ecological effects of sea otters in Oregon are most likely, what the important factors are, and, after such review, what we still don’t know about these effects.


[1] Seddon, P. J., Armstrong, D. P., and R. F. Maloney. 2007. Developing the science of reintroduction biology. Conservation Biology. 21(2): 303-312.

[2] Estes, J. A., Tinker, M. T., and J. L. Bodkin. 2009. Using ecological function to develop recovery criteria for depleted species: sea otters and kelp forests in the Aleutian Archipelago. Conservation Biology. 24(3): 852-860.

[3] Sutton, A. J., and J. P. T. Higgins. 2008. Recent developments in meta-analysis. Statistics in Medicine. 27: 625-650.

[4] Arnqvist, G., and D. Wooster. 1995. Meta-analysis: synthesizing research findings in ecology and evolution. TREE. 10(6): 236-240.

[5] Vetter, D., Rucker, G., and I. Storch. 2013. Meta-analysis: a need for well-defined usage in ecology and conservation biology. Ecosphere. 4(6): 1-13.