Throughout the world, humans rely on coastal regions for shipping and commerce, fisheries, industrial development, and increasingly for the development of marine renewable energy such as wind and wave energy . Nearshore environments, including the coastal waters of the Northern California Current (NCC), are therefore coupled social-ecological systems, at the intersection of human and biological productivity .
The NCC supports a diverse food web of ecologically and commercially important species . The nearshore region of the NCC is further shaped by a rich mosaic of complex features including rocky reefs, kelp forests, and sloping sandy bottom substrate , creating habitat for numerous species of conservation interest, including invertebrates, fish, seabirds, and marine mammals . Despite its importance, this realm poses significant challenges for vessel-based data collection, and therefore it remains relatively poorly monitored and understood.
I am excited to introduce a new project focused on these important nearshore waters, in which we will be Examining Marine mammal Ecology through Region-wide Assessment of Long-term Data (EMERALD). Since 1992, standardized surveys have been conducted between San Francisco Bay, CA, and the Columbia River, OR, to monitor the abundance of marbled murrelets, a seabird of conservation concern. Each spring and summer, researchers have simultaneously been diligently documenting the locations of harbor porpoise and gray whale sightings—two iconic marine mammal species that rely on the nearshore waters of the NCC. This rich and extensive record is rare for marine mammal data, particularly in the challenging, turbulent nearshore environment. Furthermore, harbor porpoises are cryptic, making visual sampling particularly challenging, and gray whales can be sparsely distributed, yielding low sample sizes in the absence of long-term data collection.
For the EMERALD project, we will investigate spatial and temporal distribution patterns of harbor porpoises and gray whales in relation to fluctuations in key environmental drivers. The primary goals of the project are to (1) Identify persistent hotspots in harbor porpoise and gray whale sightings over time, and (2) Examine the environmental drivers of sighting hotspots through spatial and temporal analyses.
From a first look at the data, we are already excited by some emerging patterns. In total, the dataset contains sightings of 6,763 harbor porpoise (mean 233 per year) and 530 gray whales (mean 18 per year). Preliminary data exploration reveals that harbor porpoise sightings increased in 2011-2012, predominantly between Cape Blanco, OR, and Cape Mendocino, CA. Gray whale sightings appear to follow an oscillating, cyclical pattern with peaks approximately every three years, with notable disruption of this pattern during the marine heatwave of 2014-2015. What are the drivers of sighting hotspots and spatial and temporal fluctuations in sighting rates? Time—and a quantitative analytical approach involving density estimation, timeseries analysis, and species distribution modeling—will tell.
I recently completed my PhD on the ecology and distribution of blue whales in New Zealand (for more information, see the OBSIDIAN project). Now, I am excited to apply the spatial analysis skills have been honing to a new study system and two new study species as I take on a new role in the GEMM Lab as a Postdoctoral Scholar. The EMERALD project will turn my focus to the nearshore waters close to home that I have grown to love over the past six years as a resident of coastal Oregon. The surveys I will be working with began before I was born, and I am truly fortunate to inherit such a rich dataset—a rare treat for a marine mammal biologist, and an exciting prospect for a statistical ecologist.
So, stay tuned for our findings as the project unfolds. In the meantime, I want express gratitude to Craig Strong of Crescent Coastal Research who has led the dedicated survey effort for the marbled murrelet monitoring program, without whom none of the data would exist. This project is funded by the Oregon Gray Whale License Plate funds, and we thank the gray whale license plate holders for their support of marine mammal research.
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1. Jouffray, J.-B., Blasiak, R., Norström, A. V., Österblom, H., and Nyström, M. (2020). The Blue Acceleration: The Trajectory of Human Expansion into the Ocean. One Earth 2, 43–54.
2. Sjostrom, A.J.C., Ciannelli, L., Conway, F., and Wakefield, W.W. (2021). Gathering local ecological knowledge to augment scientific and management understanding of a living coastal resource: The case of Oregon’s nearshore groundfish trawl fishery. Mar. Policy 131, 104617.
3. Bograd, S.J., Schroeder, I., Sarkar, N., Qiu, X., Sydeman, W.J., and Schwing, F.B. (2009). Phenology of coastal upwelling in the California Current. Geophys. Res. Lett. 36, 1–5.
4. Romsos, G., Goldfinger, C., Robison, R., Milstein, R., Chaytor, J., and Wakefield, W. (2007). Development of a regional seafloor surficial geologic habitat map for the continental margins of Oregon and Washington, USA. Mapp. Seafloor Habitat Charact. Geol. Assoc. Canada, Spec. Pap., 219–243.
5. Oregon Department of Fish and Wildlife (2016). Oregon Nearshore Strategy. Available at: https://oregonconservationstrategy.org/oregon-nearshore-strategy/ [Accessed January 10, 2022].
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.
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.
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. https://doi.org/10.1007/s002270000530
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. https://doi.org/10.1111/j.1748-7692.2009.00297.x
By: Alexa Kownacki, Ph.D. Student, OSU Department of Fisheries and Wildlife,
Geospatial Ecology of Marine Megafauna Lab
We live in an interesting time. Many of us academic
scientists sit in the confines of our homes, reading scientific papers,
analyzing years-worth of data, working through a years-worth of house projects,
or simply watching Netflix. While we are confined to a much smaller area,
wildlife is not.
During this challenging situation we have unique
opportunities to study what happens when people are not outside for recreation.
All of us who feel trapped inside our homes are not only saving human lives, we
are changing ecosystems. Humans are constantly molding our ecosystems on fine
and grand scales, from xeriscaping our lawns with native, drought-resistant
plants to developing large plots of land for new homes. We manipulate nature,
for better or for worse.
So, what happens when we change our behavior? Rather than
driving, we’re gardening, instead of playing at parks, we’re playing board
games at our kitchen tables; we as a society are completely changing our
habitat-use patterns. When any top predator changes its habitat-use, switches
niches, or drastically changes its behaviors, there are top-down ecosystem
effects. When one species changes its behavior, there are major downstream
impacts on predation, foraging, diet, and habitat use. For example, when
bluegill sunfish underwent large shifts in both diet and habitat, major
predator-mediated habitat use changes in other species occurred (Mittelbach
1986). There are multiple studies describing the impacts of human-mediated
drivers on ecosystems worldwide. In coastal environments, anthropogenic
activities, specifically shipping, industry, and urban development, dramatically
change both the coastal and marine ecosystems (Mead et al. 2013).
By far the most pronounced example of how an international halt on travel can alter ecosystems comes from the tragic terrorist attacks on September 11, 2001. Prior to this current, viral pandemic, the events following 9/11 were the first time that nearly all major transit stopped in the USA—including airplanes and major shipping traffic. This halt created a unique opportunity to study some of the secondary impacts, such as a reduction in shipping traffic noise, on cetaceans. Following 9/11, there was a six decibel decrease in underwater noise that co-occurred with a decrease in stress hormones of endangered North Atlantic right whales (Rolland et al. 2012). When I first read about this study, my first thought was “leave it to scientists to make the best out of a terrible situation.” Truly, learning from nature, even in the darkest of days, is an incredible skillset. Research like this inspires me to ask questions about what changes are happening in ecosystems now because of recent events. For example, the entire port of San Diego, its beaches and bays, are closed for all recreational activity and I wonder how this reduction in traffic is similar to the post-9/11 study but on bottlenose dolphins, gray whales, and pinnipeds that are coast-associated. Are urban and suburban neighborhoods slowly becoming more rural and making space for wildlife again?
Mead, A., Griffiths, C.L., Branch, G.M.,
McQuaid, C.D., Blamey, L.K., Bolton, J.J., Anderson, R.J., Dufois, F., Rouault,
M., Froneman, P.W. and Whitfield, A.K., 2013. Human-mediated drivers of
change—impacts on coastal ecosystems and marine biota of South Africa. African
Journal of Marine Science, 35(3), pp.403-425.
Mittelbach, Gary. 1986. Predator-mediated
habitat use: some consequences for species interactions. Environ Biol
Fish16, 159–169. https://doi.org/10.1007/BF00005168
Rolland, R.M., Parks, S.E., Hunt, K.E.,
Castellote, M., Corkeron, P.J., Nowacek, D.P., Wasser, S.K. and Kraus, S.D.,
2012. Evidence that ship noise increases stress in right whales. Proceedings
of the Royal Society B: Biological Sciences, 279(1737),
By Dominique Kone, Masters Student in Marine Resource Management
By now, I’m sure you’re aware of recent interests to reintroduce sea otters to Oregon. To inform this effort, my research focuses on predicting suitable sea otter habitat and investigating the potential ecological effects if sea otters are reintroduced in the future. This information will help managers gain a better understanding of the potential for sea otters to reestablish in Oregon, as well as how Oregon’s ecosystems may change via top-down processes. These analyses will address some sources of uncertainties of this effort, but there are still many more questions researchers could address to further guide this process. Here, I note some lingering questions I’ve come across in the course of conducting my research. This is not a complete list of all questions that could or should be investigated, but they represent some of the most interesting questions I have and others have in Oregon.
The questions, and our associated knowledge on each of these topics:
Is there enough available prey to support a robust sea otter population in Oregon?
Sea otters require approximately 30% of their own body weight in food every day (Costa 1978, Reidman & Estes 1990). With a large appetite, they not only need to spend most of their time foraging, but require a steady supply of prey to survive. For predators, we assume the presence of suitable habitat is a reliable proxy for prey availability (Redfern et al. 2006). Whereby, quality habitat should supply enough prey to sustain predators at higher trophic levels.
In making these habitat predictions for sea otters, we must also recognize the potential limitations of this “habitat equals prey” paradigm, in that there may be parcels of habitat where prey is unavailable or inaccessible. In Oregon, there could be unknown processes unique to our nearshore ecosystems that would support less prey for sea otters. This possibility highlights the importance of not only understanding how much suitable habitat is available for foraging sea otters, but also how much prey is available in these habitats to sustain a viable otter population in the future. Supplementing these habitat predictions with fishery-independent prey surveys is one way to address this question.
How will Oregon’s oceanographic seasonality alter or impact habitat suitability?
Sea otters along the California coast exist in an environment with persistent Giant kelp beds, moderate to low wave intensity, and year-round upwelling regimes. These environmental variables and habitat factors create productive ecosystems that provide quality sea otter habitat and a steady supply of prey; thus, supporting high densities of sea otters. This environment contrasts with the Oregon coast, which is characterized by seasonal changes in bull kelp and wave intensity. Summer months have dense kelp beds, calm surf, and strong upwellings. While winter months have little to no kelp, weak upwellings, and intense wave climates. These seasonal variations raise the question as to how these temporal fluctuations in available habitat could impact the number of sea otters able to survive in Oregon.
In Washington – an environment like Oregon – sea otters exhibit seasonal distribution patterns in response to intensifying wave climates. During calm summer months, sea otters primarily forage along the outer coast, but move into more protected areas, such as the Strait of Juan de Fuca, during winter months (Laidre et al. 2009). If sea otters were reintroduced to Oregon, we may very well observe similar seasonal movement patterns (e.g. dispersal into estuaries), but the degree to which this seasonal redistribution and reduction in foraging habitat could impact sea otter reestablishment and recovery is currently unknown.
In the event of a reintroduction, do northern or southern sea otters have a greater capacity to adapt to Oregon environments?
In the early 1970’s, Oregon’s first sea otter translocation effort failed (Jameson et al. 1982). Since then, hypotheses on the potential ecological differences between northern and southern sea otters have been proposed as potential factors of the failed effort, potentially due to different abilities to exploit specific prey species. Studies have demonstrated that northern and southern sea otters have slight morphological differences – northern otters having larger skulls and teeth than southern otters (Wilson et al. 1991). This finding has created the hypothesis that the northern otter’s larger skull and teeth allow it to consume prey with denser exoskeletons, and thereby can exploit a greater diversity of prey species. However, there appears to be a lack of evidence to suggest larger skulls and teeth translate to greater bite force. Based on morphology alone, either sub-species could be just as successful in exploiting different prey species.
A different direction to address questions around adaptability is to look at similarities in habitat and oceanographic characteristics. Sea otters exist along a gradient of habitat types (e.g. kelp forests, estuaries, soft-sediment environments) and oceanographic conditions (e.g. warm-temperature to cooler sub-Arctic waters) (Laidre et al. 2009, Lafferty et al. 2014). Yet, we currently don’t know how well or quickly otters can adapt when they expand into new habitats that differ from ones they are familiar with. Sea otters must be efficient foragers and need to acquire skills that allow them to effectively hunt specific prey species (Estes et al. 2003). Hypothetically, if we take sea otters from rocky environments where they’ve developed foraging skills to hunt sea urchins and abalones, and place them in a soft-sediment environment, how quickly would they develop new foraging skills to exploit soft-sediment prey species? Would they adapt quickly enough to meet their daily prey requirements?
In Oregon, specifically, how might climate change impact sea otters, and how might sea otters mediate climate impacts?
Climate change has been shown to directly impact many species via changes in temperature (Chen et al. 2011). Some species have specific thermal tolerances, in which they can only survive within a specified temperature range (i.e. maximum and minimum). Once the temperature moves out of that range, the species can either move with those shifting water masses, behaviorally adapt or perish (Sunday et al. 2012). It’s unclear if and how changing temperatures will impact sea otters, directly. However, sea otters could still be indirectly affected via impacts to their prey. If prey species in sea otter habitat decline due to changing temperatures, this would reduce available food for otters. Ocean acidification (OA) is another climate-induced process that could indirectly impact sea otters. By creating chemical conditions that make it difficult for species to form shells, OA could decrease the availability of some prey species, as well (Gaylord et al. 2011).
Interestingly, these pathways between sea otters and climate change become more complex when we consider the potentially mediating effects from sea otters. Aquatic plants – such as kelp and seagrass – can reduce the impacts of climate change by absorbing and taking carbon out of the water column (Krause-Jensen & Duarte 2016). This carbon sequestration can then decrease acidic conditions from OA and mediate the negative impacts to shell-forming species. When sea otters catalyze a tropic cascade, in which herbivores are reduced and aquatic plants are restored, they could increase rates of carbon sequestration. While sea otters could be an effective tool against climate impacts, it’s not clear how this predator and catalyst will balance each other out. We first need to investigate the potential magnitude – both temporal and spatial – of these two processes to make any predictions about how sea otters and climate change might interact here in Oregon.
There are several questions I’ve noted here that warrant further investigation and could be a focus for future research as this potential sea otter reintroduction effort progresses. These are by no means every question that should be addressed, but they do represent topics or themes I have come across several times in my own research or in conversations with other researchers and managers. I think it’s also important to recognize that these questions predominantly relate to the natural sciences and reflect my interest as an ecologist. The number of relevant questions that would inform this effort could grow infinitely large if we expand our disciplines to the social sciences, economics, genetics, so on and so forth. Lastly, these questions highlight the important point that there is still a lot we currently don’t know about (1) the ecology and natural behavior of sea otters, and (2) what a future with sea otters in Oregon might look like. As with any new idea, there will always be more questions than concrete answers, but we – here in the GEMM Lab – are working hard to address the most crucial ones first and provide reliable answers and information wherever we can.
Chen, I., Hill, J. K., Ohlemuller, R., Roy, D. B., and C. D. Thomas. 2011. Rapid range shifts of species associated with high levels of climate warming. Science. 333: 1024-1026.
Costa, D. P. 1978. The ecological energetics, water, and electrolyte balance of the California sea otter (Enhydra lutris). Ph.D. dissertation, University of California, Santa Cruz.
Estes, J. A., Riedman, M. L., Staedler, M. M., Tinker, M. T., and B. E. Lyon. 2003. Individual variation in prey selection by sea otters: patterns, causes and implications. Journal of Animal Ecology. 72: 144-155.
Gaylord et al. 2011. Functional impacts of ocean acidification in an ecologically critical foundation species. Journal of Experimental Biology. 214: 2586-2594.
Jameson, R. J., Kenyon, K. W., Johnson, A. M., and H. M. Wight. 1982. History and status of translocated sea otter populations in North America. Wildlife Society Bulletin. 10(2): 100-107.
Krause-Jensen, D., and C. M. Duarte. 2016. Substantial role of macroalgae in marine carbon sequestration. Nature Geoscience. 9: 737-742.
Lafferty, K. D., and M. T. Tinker. 2014. Sea otters are recolonizing southern California in fits and starts. Ecosphere.5(5).
Laidre, K. L., Jameson, R. J., Gurarie, E., Jeffries, S. J., and H. Allen. 2009. Spatial habitat use patterns of sea otters in coastal Washington. Journal of Marine Mammalogy. 90(4): 906-917.
Redfern et al. 2006. Techniques for cetacean-habitat modeling. Marine Ecology Progress Series. 310: 271-295.
Reidman, M. L. and J. A. Estes. 1990. The sea otter (Enhydra lutris): behavior, ecology, and natural history. United States Department of the Interior, Fish and Wildlife Service, Biological Report. 90: 1-126.
Sunday, J. M., Bates, A. E., and N. K. Dulvy. 2012. Thermal tolerance and the global redistribution of animals. Nature: Climate Change. 2: 686-690.
Wilson, D. E., Bogan, M. A., Brownell, R. L., Burdin, A. M., and M. K. Maminov. 1991. Geographic variation in sea otters, Ehydra lutris. Journal of Mammalogy. 72(1): 22-36.
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.
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.
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.
H1:True, dolphins are clustered and do not have a uniform distribution across this area. Spatial analysis indicated a less than a 1% likelihood that this clustered pattern could be the result of random chance (Fig. 1, z-score = -127.16, p-value < 0.0001). It is well-known that schooling fishes have a patchy distribution, which could influence the clustered distribution of their dolphin predators. In addition, bottlenose dolphins are highly social and although pods change in composition of individuals, the dolphins do usually transit, feed, and socialize in small groups.
H2:False, dolphins do not occur at higher densities in the higher latitudes of the San Diego study site. The sightings are more clumped towards the lower latitudes overall (p < 2e-16), possibly due to habitat preference. The sightings are closer to beaches with higher human densities and human-related activities near Mission Bay, CA. It should be noted, that just north of the San Diego transect is the Camp Pendleton Marine Base, which conducts frequent military exercises and could deter animals.
H3: False, during warm (positive) El Niño Southern Oscillation (ENSO) months, the dolphin sightings in San Diego were more southerly. In colder (negative) ENSO months, the dolphins were more northerly. The differences between sighting latitude and ENSO index was significant (p<0.005). Post-hoc analysis indicates that the north-south distribution of dolphin sightings was different during each ENSO state.
H4:True, dolphins are clustered around particular lagoons. Figure 5 illustrates how dolphin sightings nearest to Lagoon 6 (the San Dieguito Lagoon) are always within 0.03 decimal degrees. Because of how these data are formatted, decimal degrees is the easiest way to measure change in distance (in this case, the difference in latitude). In comparison, dolphins at Lagoon 5 (Los Penasquitos Lagoon) are distributed across distances, with the most sightings further from the lagoon.
I found a significant difference between distance to nearest lagoon in different ENSO index categories (p < 2.55e-9): there is a significant difference in distance to nearest lagoon between neutral and negative values and positive and neutral years. Therefore, I hypothesize that in neutral ENSO months compared to positive and negative ENSO months, prey distributions are changing. This is one possible hypothesis for the significant difference in lagoon preference based on the monthly ENSO index. Using a violin plot (Fig. 6), it appears that Lagoon 5, Los Penasquitos Lagoon, has the widest variation of sighting distances in all ENSO index conditions. In neutral years, Lagoon 0, the Buena Vista Lagoon has multiple sightings, when in positive and negative years it had either no sightings or a single sighting. The Buena Vista Lagoon is the most northerly lagoon, which may indicate that in neutral ENSO months, dolphin pods are more northerly in their distribution.
Takeaways to science and management:
Bottlenose dolphins have a clustered distribution which seems to be related to ENSO monthly indices, and likely, their social structures. From these data, neutral ENSO months appear to have something different happening compared to positive and negative months, that is impacting the sighting distributions of bottlenose dolphins off the San Diego coastline. More research needs to be conducted to determine what is different about neutral months and how this may impact this dolphin population. On a finer scale, the six lagoons in San Diego appear to have a spatial relationship with dolphin sightings. These lagoons may provide critical habitat for bottlenose dolphins and/or for their preferred prey either by protecting the animals or by providing nutrients. Different lagoons may have different spans of impact, that is, some lagoons may have wider outflows that create larger nutrient plumes.
Other than the Marine Mammal Protection Act and small protected zones, there are no safeguards in place for these dolphins, whose population hovers around 500 individuals. Therefore, specific coastal areas surrounding lagoons that are more vulnerable to habitat loss, habitat degradation, and/or are more frequented by dolphins, may want greater protection added at a local, state, or federal level. For example, the Batiquitos and San Dieguito Lagoons already contain some Marine Conservation Areas with No-Take Zones within their reach. The city of San Diego and the state of California need better ways to assess the coastlines in their jurisdictions and how protecting the marine, estuarine, and terrestrial environments near and encompassing the coastlines impacts the greater ecosystem.
This dive into my data was an excellent lesson in spatial scaling with regards to parsing down my data to a single study site and in matching my existing data sets to other data that could help answer my hypotheses. Originally, I underestimated the robustness of my data. At first, I hesitated when considering reducing the dolphin sighting data to only include San Diego because I was concerned that I would not be able to do the statistical analyses. However, these concerns were unfounded. My results are strongly significant and provide great insight into my questions about my data. Now, I can further apply these preliminary results and explore both finer and broader scale resolutions, such as using the more precise ENSO index values and finding ways to compare offshore bottlenose dolphin sighting distributions.
I have the privilege of studying the largest animals on the planet: blue whales (Balaenoptera musculus). However, in order to understand the ecology, distribution, and habitat use patterns of these ocean giants, I have dedicated the past several months to studying something much smaller: krill (Nyctiphanes australis). New Zealand’s South Taranaki Bight region (“STB”, Figure 1) is an important foraging ground for a unique population of blue whales [1,2]. A wind-driven upwelling system off of Kahurangi Point (the “X” in Figure 1) generates productivity in the region , leading to an abundance of krill , the desired blue whale prey .
What happened to the blue whales’ food source under these different conditions in 2016? Before I share some preliminary findings from my recent analyses, it is important to note that there are many possible ways to measure krill availability. For example, the number of krill aggregations, as well as how deep, thick, and dense those aggregations are in an area will all factor into how “desirable” krill patches are to a blue whale. While there may not be “more” or “less” krill from one year to the next, it may be more or less accessible to a blue whale due to energetic costs of capturing it. Here is a taste of what I’ve found so far:
In 2016, when surface waters were warm, the krill aggregations were significantly deeper than in the “typical” years (ANOVA, F=7.94, p <0.001):
The number of aggregations was not significantly different between years, but as you can see in the plot below (Figure 4) the krill were distributed differently in space:
While the bulk of the krill aggregations were located north of Cape Farewell under typical conditions (2014 and 2017), in the warm year (2016) the krill were not in this area. Rather, the area with the most aggregations was offshore, in the western portion of our study region. Now, take a look at the same figure, overlaid with our blue whale sighting locations:
Where did we find the whales? In each year, most whale encounters were in the locations where the most krill aggregations were found! Not only that, but in 2016 the whales responded to the difference in krill distribution by shifting their distribution patterns so that they were virtually absent north of Cape Farewell, where most sightings were made in the typical years.
The above figures demonstrate the importance of studying an ecosystem. We could puzzle and speculate over why the blue whales were further west in the warm year, but the story that is emerging in the krill data may be a key link in our understanding of how the ecosystem responds to warm conditions. While the focus of my dissertation research is blue whales, they do not live in isolation. It is through understanding the ecosystem-scale story that we can better understand blue whale ecology in the STB. As I continue modeling the relationships between oceanography, krill, and blue whales in warm and typical years, we are beginning to scratch the surface of how blue whales may be responding to their environment.
Torres LG. 2013 Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zeal. J. Mar. Freshw. Res.47, 235–248. (doi:10.1080/00288330.2013.773919)
Barlow DR et al. 2018 Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger. Species Res.36, 27–40. (doi:https://doi.org/10.3354/esr00891)
Shirtcliffe TGL, Moore MI, Cole AG, Viner AB, Baldwin R, Chapman B. 1990 Dynamics of the Cape Farewell upwelling plume, New Zealand. New Zeal. J. Mar. Freshw. Res.24, 555–568. (doi:10.1080/00288330.1990.9516446)
Bradford-Grieve JM, Murdoch RC, Chapman BE. 1993 Composition of macrozooplankton assemblages associated with the formation and decay of pulses within an upwelling plume in greater cook strait, New Zealand. New Zeal. J. Mar. Freshw. Res.27, 1–22. (doi:10.1080/00288330.1993.9516541)
Gill P. 2002 A blue whale (Balaenoptera musculus) feeding ground in a southern Australian coastal upwelling zone. J. Cetacean Res. Manag.4, 179–184.
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. 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.
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. 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.
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. 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. 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. 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.
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.
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.
 Seddon, P. J., Armstrong, D. P., and R. F. Maloney. 2007. Developing the science of reintroduction biology. Conservation Biology. 21(2): 303-312.
 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.
 Sutton, A. J., and J. P. T. Higgins. 2008. Recent developments in meta-analysis. Statistics in Medicine. 27: 625-650.
 Arnqvist, G., and D. Wooster. 1995. Meta-analysis: synthesizing research findings in ecology and evolution. TREE. 10(6): 236-240.
 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.
By Dominique Kone, Masters Student in Marine Resource Management
Species reintroductions can be hotly contested issues because they can negatively impact other species, ecosystems, and society, as well as failing, altogether. The uncertainty of their outcomes forces stakeholder groups to form their own opinions on whether it’s a good idea to proceed with a reintroduction. When you have several groups with conflicting values and views, managers need to focus on the information most important for them to make a well-informed decision on whether to pursue a reintroduction.
As researchers, we can play an important role by carefully considering and addressing these views through our research, if the appropriate data is available. Despite being in the early days of our study on the potential sea otter reintroduction to Oregon, we have already heard several perspectives regarding its potential success, the type of research we should do, and if sea otters should be brought back to Oregon. Here, I present some of the most interesting and relevant opinions, perspectives, and theories I’ve heard regarding this reintroduction idea.
The first reintroduction failed because of X, Y, and Z.
From 1970-1971, managers translocated 93 sea otters to Oregon in a reintroduction effort (Jameson et al. 1982). However, in a matter of 5-6 years, all sea otters disappeared, and the effort was considered a failure. Researchers have theorized that sea otters left Oregon due to a lack of suitable habitat and prey, or to return home to sites from which they were captured. Others have reasoned that managers should have introduced southern sea otters instead of northern sea otters, suggesting one subspecies’ genetic pre-disposition may improve their chance for survival.
Knowing the reasons for this failure may help managers avoid these causes in a future reintroduction attempt and increase its chance of success. We, as scientists, can also gain insight from knowing these causes because this may help us better tailor our research to potentially investigate whether those causes still pose a threat to sea otters during a second attempt. Unfortunately, we lack concrete evidence on what exactly caused this failure, but we can still work to test some these theories.
An otter is an otter, no matter where you put it.
There is evidence that northern and southern sea otters are genetically distinct, to a certain degree (Valentine et al. 2008, Larson et al. 2012), and hypotheses have been put forward that the two subspecies may be behaviorally- and ecologically-distinct, too. Studies have shown that northern and southern sea otters have different sized and shaped skulls and teeth, which researchers hypothesize may be a specialized foraging adaptation for consuming different prey species (Campbell & Santana 2017, Timm-Davis et al. 2015). This view suggests that each subspecies has developed unique traits to adapt to the environmental conditions specific to their current ranges. Therefore, when considering which subspecies to bring to Oregon, managers should reintroduce the subspecies with traits better-suited to cope with the types of habitat, prey assemblages, and oceanographic conditions specific to Oregon.
However, other scientists hold the opposite view, and argue that “an otter is an otter” no matter where you put it. This perspective suggests that both subspecies have an equal chance at surviving in any type of suitable habitat because all otters behave in similar ways. Therefore, ecologically, it may not matter which subspecies managers bring to Oregon.
Oregon doesn’t have enough sea otter habitat.
Kelp is considered important sea otter habitat. In areas with high sea otter densities, such as central and southern California, kelp forests are persistent throughout the year. However, in Oregon, our kelp primarily consists of bull kelp – a slightly more fragile species compared to the durable giant kelp in California. In winter, this bull kelp gets dislodged during intense storms, resulting in seasonal changes in kelp availability. Managers worry that this seasonality could reduce the amount of suitable habitat, to the point where Oregon may not be able to support sea otters.
Yet, we know sea otters used to exist here; therefore, we can assume there must have been some suitable habitat that may persist today. Furthermore, sea otters use a range of habitats, including estuaries, bays, and reefs (Laidre et al. 2009, Lafferty & Tinker 2014, Kvitek et al. 1988). Therefore, even during times when kelp is less abundant, sea otters could use these other forms of habitat along the Oregon coast. Luckily, we have the spatial tools and data to assess how much, where, and when we have suitable habitat, and I will specifically address this in my thesis.
They’ll eat everything!
Sea otters are famous for their voracious appetites for benthic invertebrates, some of which are of commercial and recreational importance to nearshore fisheries. In some cases, sea otters have significantly reduced prey densities, such as sea urchins and Dungeness crab (Garshelis & Garshelis 1984, Estes & Palmisano 1974). However, without a formal analysis, it’s difficult to know if sea otters will have similar impacts on Oregon’s nearshore species, as well as at spatial scale these impacts will occur and whether our fisheries will be affected. We can predict where sea otters are likely to occur based on the presence of suitable habitat, but foraging impacts could be more localized or widespread across sea otter’s entire potential range. To better anticipate these impacts, managers will need an understanding of how much sea otters eat, where foraging could occur based on the availability of prey, and where sea otters and fisheries are likely to interact. I will also address this concern in my thesis.
To reintroduce or not to reintroduce? That is the question.
I have found that many scientists and managers have strong opinions on whether it’s appropriate to bring sea otters back to Oregon. Those who argue against a reintroduction often highlight many of the theories already mentioned here – lack of habitat, potential impacts to fisheries, and genetics. While other opponents provided more logistical and practical justifications, such as confounding politics, as well as difficulties in getting public support and regulatory permission to move a federally-listed species.
In contrast, proponents of this idea argue that a reintroduction could augment the recovery of the species by providing additional habitat for the species to rebound to pre-exploitation levels, as well as allowing for increased gene flow between southern and northern sea otter populations. Other proponents have brought up potential benefits to humans, such restoring ecosystem services, providing an economic boost through tourism, or preserving tribal and cultural connections. Such benefits may be worth attempting another reintroduction effort.
As you can see, there are several opinions and perspectives related to a potential sea otter reintroduction to Oregon. While it’s important to consider all opinions, managers still need facts to make key decisions. Scientists can play an important role in providing this information, so managers can make a well-informed decision. Oregon managers have not yet decided whether to proceed with a sea otter reintroduction, but our lab is working to provide them with reliable and accurate science, so they may form their own opinions and arrive at their own decision.
Estes, J. A. and J. F. Palmisano. 1974. Sea otters: the role in structuring nearshore communities. Science. 185: 1058-1060.
Garshelis, D. L. and J. A. Garshelis. 1984. Movements and management of sea otters in Alaska. The Journal of Wildlife Management. 48: 665-678.
Jameson, R. J, Kenyon, K. W., Johnson, A. M., and H. M. Wight. 1982. History and status of translocated sea otter populations in North America. Wildlife Society Bulletin. 10: 100-107.
Lafferty, K. D., and M. T. Tinker. 2014. Sea otters are recolonizing southern California in fits and starts. Ecosphere. 5(5).
Laidre, K. L., Jameson, R. J., Gurarie, E., Jeffries, S. J., and H. Allen. 2009. Spatial habitat use patterns of sea otters in coastal Washington. Journal of Mammalogy. 90(4): 906-917.
Kvitek, R. G. ,Fukayama, A. K., Anderson, B. S., and B. K. Grimm. 1988. Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon. Marine Biology. 98: 157-167.
Larson, S., Jameson, R., Etnier, M., Jones, T., and R. Hall. 2012. Genetic diversity and population parameters of sea otters, Enhydra lutris, before fur trade extirpation from 1741-1911. PLoS ONE. 7(3).
Timm-Davis, L. L, DeWitt, T. J., and C. D. Marshall. 2015. Divergent skull morphology supports two trophic specializations in otters (Lutrinae). PLoS ONE. 10(12).
Valentine et al. 2008. Ancient DNA reveals genotypic relationships among Oregon populations of the sea otter (Enhydra lutris). Conservation Genetics. 9:933-938.
By Dawn Barlow, MSc student, Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab
In 2013, Leigh first published a hypothesis that the South Taranaki Bight region between New Zealand’s North and South Islands is important habitat for blue whales (Torres 2013). Since then, we have collected three years of data and conducted dedicated analyses, so we now understand that a unique population of blue whales is found in New Zealand, and that they are present in the South Taranaki Bight year-round (Barlow et al. in press).
This research has garnered quite a bit of political and media attention. A major platform item for the New Zealand Green Party around the last election was the establishment of a marine mammal sanctuary in the South Taranaki Bight. When the world’s largest seismic survey vessel began surveying the South Taranaki Bight this summer for more oil and gas reserves using tremendously loud airguns, there were rallies on the lawn in front of Parliament featuring a large inflatable blue whale that the protesters affectionately refer to as “Janet”. Needless to say, blue whales have made their way into the spotlight in New Zealand.
Now that we know there is a unique population of blue whales in New Zealand, what is next? What’s next for me is an exciting combination of both ecology and conservation. If an effective sanctuary is to be implemented, it needs to be more than a simple box drawn on a map to check off a political agenda item—the sanctuary should be informed by our best ecological knowledge of the blue whales and their habitat.
In July, Leigh and I will attend the Society for Conservation Biology meeting in Wellington, New Zealand, and I’ll be giving a presentation titled “Cloudy with a chance of whales: Forecasting blue whale presence based on tiered, bottom-up models”. I’ll be the first to admit, I am not yet forecasting blue whale presence. But I am working my way there, step-by-step, through this tiered, bottom-up approach. In cetacean habitat modeling, we often assume that whale distribution on a foraging ground is determined by their prey’s distribution, and that satellite images of temperature and chlorophyll-a provide an accurate picture of what is going on below the surface. Is this true? With our three years of data including in situ oceanography, krill hydroacoustics, and blue whale distribution and behavior, we are in a unique position to test some of those assumptions, as well as provide managers with an informed management tool to predict blue whale distribution.
What questions will we ask using our data? Firstly, can in situ oceanography (i.e., thermocline depth and temperature, mixed layer depth) predict the distribution and density of blue whale prey (krill)? Then, can those prey patterns be accurately predicted in the absence of oceanographic measurements, using just satellite images? Next, we’ll bring the blue whales back into the picture to ask: can we predict blue whale distribution based on our in situ measurements of oceanography and prey? And finally, in the absence of in situ measurements (which is most often the case), can we forecast where the whales will be based just on remotely-sensed images of the region?
So, cloudy with a chance of whales? Well, you’ll have to stay tuned for that story in the coming months. In the meantime, I can tell you that as daunting as it is to aggregate so many data streams, each step of the way has a piece of the story to tell. I can’t wait to see how it falls together, both from an ecological modeling perspective and a conservation management objective.
Torres, L. G. (2013). Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zealand Journal of Marine and Freshwater Research, 47(2), 235-248.
Barlow, D. R., Torres, L. G., Hodge, K. B., Steel, D. Baker, C. S., Chandler, T. E., Bott, N., Constantine, R., Double, M. C., Gill, P., Glasgow, D., Hamner, R. M., Lilley, C., Ogle, M., Olson, P. A., Peters, C., Stockin, K. A., Tessaglia-Hymes, C. T., Klinck, H. (in press). Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endangered Species Research.
By Dominique Kone, Masters Student in Marine Resource Management
When considering a species reintroduction into an area, it is important to assess the suitability of the area’s habitat before such efforts begin. By doing this assessment at the outset, managers and conservationists can gain a better understanding of the capacity of the area to support a viable population overtime, and ultimately the success of the reintroduction. However, to do a habitat assessment, researchers must first have a base understanding of the species’ ecological characteristics, behavior, and the physical habitat features necessary for the species’ survival. For my thesis, I plan to conduct a similar assessment to identify suitable sea otter habitat to inform a potential sea otter reintroduction to the Oregon coast.
To start my assessment, I conducted a literature review of studies that observed and recorded the various types of habitats where sea otters currently exist. In my research, I learned that sea otters use in a range of environments, each with a unique set of habitat characteristics. With so many features to sort through, I have focused on specific habitat features that are consistent across most of the current range of sea otters – from Alaska to California – and are important for at least some aspects of sea otters’ everyday life or behavior, specifically foraging. Focusing my analysis on foraging habitat makes sense as sea otters require around 30% of their body weight in food every day (Costa 1978, Reidman & Estes 1990). Meaning sea otters spend most of their day searching for food.
Here, I present four habitat features I will incorporate into my analysis and explain why these features are important for sea otter foraging behavior and survival.
Kelp: Sea otters are famously known for the benefits they provide to kelp forests. In the classic three-trophic-level model, sea otters allow for the growth of kelp by keeping sea urchins – consumers of kelp – in check (Estes & Palmisano 1974). Additionally, sea otters and kelp have a mutually-beneficial relationship. Sea otters will often wrap themselves amongst the top of kelp stocks while feeding, resting, or grooming to prevent being carried away by surface currents. Meanwhile, it’s thought that kelp provide a refuge for sea otters seeking to avoid predators, such as sharks, as well as their prey.
Distance from Kelp: The use of kelp, by sea otters, is relatively straight-forward. Yet, kelp can still have an influence on sea otter behavior even when not used directly. A 2014 study found that sea otters along the southern California coast were almost 10 times more likely to be located within kelp habitat than outside, while outside kelp beds sea otter numbers declined with distance from the edge of kelp canopies. Sea otters will often forage outside or next to kelp canopies when prey’s available, and even sometimes to socialize in age- or sex-specific rafts (Lafferty & Tinker 2014). These findings indicate that sea otters can and do regularly disperse away from kelp habitat, but because they’re so dependent on kelp, they don’t stray very far.
Seafloor Substrate: Sea otters forage over a variety of sediment substrates, including rocks, gravel, seagrass, and even sometimes sand. For example, sea otters hunt sea urchins over rocky substrates, while in other areas they may hunt for crabs in seagrass beds (Estes & Palmisano 1974, Hughes et al. 2014). The type of substrate sea otters forage in typically depends on the substrate needs of their target prey species. Despite some variability across their range, sea otters predominantly forage in rocky substrate environments. Rocky substrate is also necessary for kelp, whose holdfasts need to attach to hard, stable surfaces (Carney et al. 2005).
Depth: Seafloor depth plays a pivotal role in sea otter foraging behavior and therefore acts as a natural boundary that determines how far away from shore sea otters distribute. Many of the prey species sea otters eat – including sea urchins, crabs, and snails – live on the seafloor of the inner continental shelf, requiring sea otters to dive when foraging. Interestingly, sea otters exhibit a non-linear relationship with depth, where most individuals forage at intermediate depths as opposed to extremely shallow or deep waters. One study found the average foraging depth to be around 15 meters (Lafferty & Tinker 2014). This behavior results in a hump-shaped distribution of diving patterns as illustrated in Figure 1 below.
Of course, local conditions and available habitat are always a factor. For example, a study found that sea otters along the coast of Washington foraged further from shore and in slightly shallower environments than sea otters in California (Laidre et al. 2009), indicating that local topography is important in determining distribution. Additionally, diving requires energy and limits how deep sea otters are able to forage for prey. Therefore, diving patterns are not only a function of local topography, but also availability of prey and foraging efficiency in exploiting that prey. Regardless, most sea otter populations follow this hump-shaped diving pattern.
These features are not a complete list of all habitat characteristics that support viable sea otter populations, but seem to be the most consistent throughout their entire range, as well as present in Oregon’s nearshore environment – making them ideal features to include in my analysis. Furthermore, other studies that have predicted suitable sea otter habitat (Tinker et al. 2017), estimated carrying capacity as a product of suitable habitat identification (Laidre et al. 2002), or simply observed sea otter foraging behavior (Estes & Palmisano 1974), have echoed the importance of these four habitat features to sea otter survival.
As with most reintroduction efforts, the process of identifying suitable habitat for the species of interest can be complicated. No two ecosystems or habitats are exactly alike and each comprise their own unique set of physical features and are impacted by environmental processes to varying degrees. The Oregon coast consists of a unique combination of oceanographic conditions and drivers that likely impact the degree and amount of available habitat to sea otters. Despite this, by focusing on the habitat features that are consistently preferred by sea otters across most of their range, I will be able to identify habitat most suitable for sea otter survival in Oregon. The questions of where this habitat is and how much is available are what I’ll determine soon, so stay tuned.
Carney, L. T., Robert Waaland, J., Kilinger, T., and K. Ewing. 2005. Restoration of the bull kelp Nereocystis luetkeana in nearshore rocky habitats. Marine Ecology Progress Series. 302: 49-61.
Costas, D. P. 1978. The ecological energetics, waters, and electrolyte balance of the California sea otter (Enhydra lutris). Ph.D. dissertation, University of California, Santa Cruz.
Estes, J. A. and J. F. Palmisano. 1974. Sea otters: their role in structuring nearshore communities. Science. 185(4156): 1058-1060.
Hughes et al. 2014. Recovery of a top predator mediate negative eutrophic effects on seagrass. Proceedings of the National Academy of Sciences. 110(38): 15313-15318.
Lafferty, K. D. and M. T. Tinker. 2014. Sea otters are recolonizing southern California in fits and starts. Ecosphere. 5(5): 1-11.
Laidre et al. 2002. Estimates of carrying capacity for sea otters in Washington state. Wildlife Society Bulletin. 30(4): 1172-1181.
Laidre et al. 2009. Spatial habitat use patterns of sea otters in coastal Washington. Journal of Mammalogy. 90(4): 906-917.
Tinker et al. 2017. Southern sea otter range expansion and habitat use in the Santa Barbara Channel, California: U.S. Geological Survey Open-File Report 2017-1001 (OCS Study BOEM 2017-022), 76 p., http://doi.org/10.3133/ofr20171001.
Reidman, M. L. and J. A. Estes. 1990. The sea otter (Enhydra lutris): behavior, ecology, and natural history. United States Department of the Interior, Fish and Wildlife Service, Biological Report. 90: 1-126.