Geology topics
James L. Bodkin
Publications and source records attributed to James L. Bodkin.
Sea otter-mediated linkages among oceanic, nearshore, and terrestrial ecosystems
Seemingly distinct ecological communities are often linked by both direct and indirect interactions that can be difficult to discern. A widely recognized interecosystem linkage is the transfer of marine-derived nutrients to terrestrial systems via anadromous fish. In this chapter, we explore other linkages that move marine-derived nutrients between ocean and terrestrial ecosystems, focusing on the role of nearshore marine ecosystems . Through oceanic nutrients and biomass that originate from phytoplankton, and nearshore nutrients and biomass that derive mainly from macroalgae, there are multiple pathways to subsidize terrestrial ecosystems. We discuss various pathways identified through multiple independent studies and explore how those subsidies may be related to sea otter presence. Recovery of sea otters and subsequent changes to nearshore communities, for example, changes in kelp, invertebrate and vertebrate abundance, and diversity, may in turn influence coastal terrestrial ecosystems and species including bears, wolves, foxes, river otters, shorebirds, and other consumers. As offshore oceanic productivity declines globally, we ask how the return of the sea otter reinvigorates diminished and lost linkages between marine and terrestrial environments through direct predator–prey linkages, and indirectly through the enhancement of the nearshore marine environment.
Planktonic to sessile: Drivers of spatial and temporal variability across barnacle life stages and indirect effects of the Pacific Marine Heatwave
Barnacles are a foundation species in intertidal habitats. During the Pacific Marine Heatwave (PMH), intertidal barnacle cover increased in the northern Gulf of Alaska (GoA); however, the role of pelagic larval supply in this increase was unknown. Using long-term monitoring data on intertidal benthic (percent cover) and pelagic larval populations (nauplii and cyprid concentrations), we examined potential environmental drivers (temperature, chlorophyll-a, mixed layer depth) of larval concentration and whether including larval concentration at regional and annual scales improved intertidal barnacle percent cover models in two study regions in the GoA. In both regions, larval concentrations were slightly higher following the PMH. Percent cover models were improved by including cyprid concentrations (but not nauplii), and the effect strength varied by site and tidal elevation. This indicates that larval concentration contributes as a bottom–up driver of benthic barnacle abundance. There is little evidence of a direct effect of the PMH on either life stage. Instead, our results may illustrate the positive feedback between life stages, where higher adult benthic abundance increased larval concentrations, which then supplied more new recruits to the benthos. As heatwaves continue to occur, integrating various data types can provide insights into factors influencing both benthic and pelagic communities.
Relative importance of macroalgae and phytoplankton to nearshore consumers and growth across climatic conditions in the northern Gulf of Alaska
Macroalgae and phytoplankton support the base of highly productive nearshore ecosystems in cold-temperate regions. To better understand their relative importance to nearshore food webs, this study considered four regions in the northern Gulf of Alaska where three indicator consumers were collected, filter-feeding mussels ( Mytilus trossulus ), pelagic-feeding Black Rockfish ( Sebastes melanops ), and benthic-feeding Kelp Greenling ( Hexagrammos decagrammus ). The study objectives were to (1) estimate the proportional contributions of macroalgal and phytoplankton organic matter using carbon and nitrogen stable isotopes, (2) determine if macroalgal use affected consumer growth using annual growth rings in shells or otoliths, and (3) describe changes in organic matter use and growth during the Pacific Marine Heatwave (PMH; 2014–2016) in one consumer, mussels. Macroalgae were the major organic matter source (> 60%) to the diet for all three consumers. The relationships between macroalgal contribution and growth were neutral for both fish species and significantly positive for mussels. During the PMH, mussels had a drop (> 10%) in macroalgal contributions and grew 45% less than in other time periods. Simultaneously, the relationship between macroalgal contribution and mussel growth was strongest during the PMH, explaining 48% variation compared to 3–12% before or after the PMH. Collectively, the results suggest that macroalgae is likely more important to cold-temperate nearshore food webs than phytoplankton. Management actions aimed at conserving and expanding macroalgae are likely to benefit nearshore food webs under all climate scenarios and especially during marine heatwaves.
Lack of strong responses to the Pacific marine heatwave by benthivorous marine birds indicates importance of trophic drivers
The Pacific marine heatwave (PMH) of 2014-2016 was an intense, long-lasting environmental disturbance expressed throughout the north Pacific. While dramatic consequences of the PMH on pelagic food webs have been well documented, effects on nearshore food webs, i.e., those based on macroalgae primary productivity, benthic invertebrate intermediate consumers, and specialized benthivorous top predators including some marine birds, are not well understood. We conducted summer and winter coastline surveys in two National Parks in the northern Gulf of Alaska from 2006 – 2022. We evaluated changes in abundance of benthivorous marine birds in relation to the heatwave, after accounting for effects of season and region. We also evaluated changes in abundance of nearshore benthic invertebrate prey data to allow specific consideration of a prey-based mechanism for effects of the PMH across food webs. We found that benthivorous marine birds, consisting largely of sea ducks and shorebirds, did not show a strong response to the PMH, unlike significant effects demonstrated by piscivorous birds in pelagic biomes. Unlike extreme reductions in quantity and quality of forage fish documented in other studies, we found that common benthic invertebrate prey abundance remained relatively stable, with only minor increases or decreases, in association with the PMH. Our results support the hypothesis that food availability has a strong mediating effect of the PMH on upper trophic levels across food webs. These findings show how a large-scale environmental perturbation affects biological communities through trophic pathways, provides insight into ecosystem resiliency, and can inform management strategies in the face of persistent climate change.
Gene expression and wildlife health: Varied interpretations based on perspective
We evaluated wildlife population health from the perspective of statistical means vs. variances. We outlined the choices necessary to provide the framework for our study. These consisted of spatial and temporal boundaries (e.g., choice of sentinel species, populations, time frame), measurement techniques (molecular to population level), and appropriate statistical analyses. We chose to assess the health of 19 sea otter populations, located in the north Pacific from the Aleutian Islands, AK, to Santa Barbara, CA, and varying in population growth rates and length of occupancy. Our focal metric was gene expression (i.e., mRNA transcripts) data that we had previously generated across sea otter populations as a measure of population health. We used statistical methods with different approaches (i.e., means vs. variances) and examined the subsequent interpretive outcomes and how these influence our assessment of “health.” Interpretations based on analyses using variances versus means overlapped to some degree. In general, sea otter populations with low variation in gene expression were limited by food resources and at or near carrying capacity. In populations where the variation in gene expression was moderate or high, four out of five populations were increasing in abundance, or had been recently increasing. Where we had additional information on sources of stressors at the level of the population, we were able to draw inferences from those stressors to specific gene expression results. For example, gene expression patterns of sea otters from Western Prince William Sound were consistent with long term exposure to petroleum hydrocarbons, whereas in Kachemak Bay, patterns were consistent with exposure to algal toxins. Ultimately, determination of population or ecosystem health will be most informative when multiple metrics are examined across disciplines in the context of specific scenarios and goals.
Revealing the extent of sea otter impacts on bivalve prey through multi-trophic monitoring and mechanistic models
Sea otters are apex predators that can exert considerable influence over the nearshore communities they occupy. Since facing near extinction in the early 1900s, sea otters are making a remarkable recovery in Southeast Alaska, particularly in Glacier Bay, the largest protected tidewater glacier fjord in the world. The expansion of sea otters across Glacier Bay offers both a challenge to monitoring and stewardship and an unprecedented opportunity to study the top-down effect of a novel apex predator across a diverse and productive ecosystem. Our goal was to integrate monitoring data across trophic levels, space, and time to quantify and map the predator–prey interaction between sea otters and butter clams Saxidomus gigantea , one of the dominant large bivalves in Glacier Bay and a favoured prey of sea otters. We developed a spatially-referenced mechanistic differential equation model of butter clam dynamics that combined both environmental drivers of local population growth and estimates of otter abundance from aerial survey data. We embedded this model in a Bayesian statistical framework and fit it to clam survey data from 43 intertidal and subtidal sites across Glacier Bay. Prior to substantial sea otter expansion, we found that butter clam density was structured by an environmental gradient driven by distance from glacier (represented by latitude) and a quadratic effect of current speed. Estimates of sea otter attack rate revealed spatial heterogeneity in sea otter impacts and a negative relationship with local shoreline complexity. Sea otter exploitation of productive butter clam habitat substantially reduced the abundance and altered the distribution of butter clams across Glacier Bay, with potential cascading consequences for nearshore community structure and function. Spatial variation in estimated sea otter predation processes further suggests that community context and local environmental conditions mediate the top-down influence of sea otters on a given prey. Overall, our framework provides high-resolution insights about the interaction among components of this food web and could be applied to a variety of other systems involving invasive species, epidemiology or migration.
Brown bear–sea otter interactions along the Katmai coast: Terrestrial and nearshore communities linked by predation
Sea otters were extirpated throughout much of their range by the maritime fur trade in the 18th and 19th centuries, including the coast of Katmai National Park and Preserve in southcentral Alaska. Brown bears are an important component of the Katmai ecosystem where they are the focus of a thriving ecotourism bear-viewing industry as they forage in sedge meadows and dig clams in the extensive tidal flats that exist there. Sea otters began reoccupying Katmai in the 1970s where their use of intertidal clam resources overlapped that of brown bears. By 2008, the Katmai sea otter population had grown to an estimated 7,000 animals and was likely near carrying capacity; however, in 2006–2015, the age-at-death distribution (AADD) of sea otter carcasses collected at Katmai included a higher-than-expected proportion of prime-age animals compared to most other sea otter populations in Alaska. The unusual AADD warranted scientific investigation, particularly because the Katmai population is part of the Threatened southwest sea otter stock. Brown bears in Katmai are known to prey on marine mammals and sea otters, but depredation rates are unknown; thus, we investigated carnivore predation, especially by brown bears, as a potential explanation for abnormally high prime-age otter mortality. We installed camera traps at two island-based marine mammal haulout sites within Katmai to gather direct evidence that brown bears prey on seals and sea otters. Over a period of two summers, we gathered photo evidence of brown bears making 22 attempts to prey on sea otters of which nine (41%) were successful and 12 attempts to prey on harbor seals of which one (8%) was successful. We also developed a population model based on the AADD to determine if the living population is declining, as suggested by the high proportion of prime-age animals in the AADD. We found that the population trend predicted by the modeled AADDs was contradictory to aerial population surveys that indicated the population was not in steep decline but was consistent with otter predation. Future work should focus on the direct and indirect effects these top-level predators have on each other and the coastal community that connects them.
Where land and sea meet: Brown bears and sea otters
In Katmai National Park, Alaska, USA, we have seen changes in the number of brown bears and sea otters. The number of animals of a species a habitat can support is called carrying capacity. Even though bears live on land and sea otters live in the ocean, these two mammals share coastal habitats. Bears eat salmon, other fish, plants, clams, and beached whales. Sea otters feed on clams and other marine invertebrates. All these foods are influenced by the ocean. Recently, we have seen fewer bears but more sea otters! What changed? Many things, but several observations point to the ocean. There are fewer salmon, whales, and clams, so bears rely more on plants for food. Fewer clams mean sea otters must work harder to find food. Our studies are helping us to understand how and why carrying capacity for a given species may change over time.
Evidence of increased mussel abundance related to the Pacific marine heatwave and sea star wasting
Mussels occupy a key middle trophic position in nearshore food webs linking primary producers to predators. Climate-related environmental changes may synergistically combine with changes in predator abundance to affect intertidal ecosystems. We examined the influence of two major events on mussel ( Mytilus trossulus ) abundance in the northern Gulf of Alaska: the recent Pacific marine heatwave (PMH, 2014–2016) and an outbreak of sea star wasting (SSW). We investigated how mussel abundance changed since the onset of SSW and whether the density of predatory sea stars or PMH-related temperature metrics explain variation in mussel abundance. Sea stars and mussels were surveyed since 2005 approximately annually in four regions of the northern Gulf of Alaska: Katmai (KATM), Kachemak Bay (KBAY), Kenai Fjords (KEFJ) and western Prince William Sound (WPWS). Mussel percent cover in the mid-intertidal increased 1–3 years after declines in sea stars at all regions and in the low-intertidal at KATM, KBAY, and KEFJ, but not at WPWS. After the onset of SSW, large (≥20 mm length) mussel density and mussel bed width increased at KATM but not the other regions. Total mussel densities, including recruits, did not differ before and after the onset of SSW. The total number of sea stars significantly explained variation in mussel metrics, but the proportions of the three sea star species examined did not. We did not find strong evidence for direct effects of temperature on mussels. The effects of the PMH and the SSW outbreak appear to have combined, with increased temperatures indirectly benefiting mussels in concert with relaxed top-down pressure from sea stars, allowing for increased mussel abundance. Changing mussel abundance may affect intertidal local productivity and the abundance or performance of other nearshore consumers of mussels.
Sea otter population collapse in southwest Alaska: Assessing ecological covariates, consequences, and causal factors
Sea otter ( Enhydra lutris ) populations in southwest Alaska declined substantially between about 1990 and the most recent set of surveys in 2015. Here we report changes in the distribution and abundance of sea otters, and covarying patterns in reproduction, mortality, body size and condition, diet and foraging behavior, food availability, health profiles, and exposure to environmental contaminants over this 25-yr period. The population decline, which resulted in densities on the order of 5% of environmental carrying capacity, ranged from Attu Island in the west to about Castle Cape (on the south side of the Alaska Peninsula) in the east. Remaining sea otters moved closer to shore and into shallow, protected habitats. Reproductive rates appeared unchanged with the decline. Although the demographic cause of the decline was clearly elevated mortality, stranded carcasses were rare or absent. The net rate of energy gain by foraging sea otters, body length and condition, and prey biomass density, all increased after the decline and varied inversely with sea otter population density beyond the area of decline. Sea otters within the area of decline showed no increases in health anomalies, disease, contaminant exposure, or abnormal gene transcription patterns as compared to animals outside the area of decline. These collective findings are inconsistent with nutritional limitation, disease, or environmental contaminants, and consistent with predation (or possibly some other density-independent factor) as the reason for the sea otter population decline. Our approach and analyses provide a broad conceptual template for thinking about and assessing the causes of wildlife population declines.
Sea otter foraging behavior
Sea otters are marine specialists but diet generalists, which feed primarily on benthic mega-invertebrates (i.e., body dimension >1 cm). They locate and capture epibenthic and infaunal prey with their forepaws by relying on vision and tactile sensitivity during short-duration dives (generally <2 min) in shallow waters (routine dives <30 m and maximum dive depth ~100 m) of the littoral zone. Sea otters have an elevated resting metabolic rate and small or no energy reserves in the form of blubber, so they feed every 3–4 h. Foraging dives often occur in bouts (i.e., two or more consecutive dives), which may last several hours with 1–2 min between dives, depending on the type of prey. Sea otters consume small or soft prey entirely or use their teeth or stone tools to access the flesh of mega-invertebrates with a shell, test, or exoskeleton. The daily percentage of time that sea otters devote to foraging depends on age, sex, presence of a pup, time of year, and prey abundance, which varies geographically, seasonally, and episodically. In areas occupied by sea otters for many years, epifaunal prey generally decline first followed by infaunal species, and this may result in greater foraging effort and diet specialization associated with density-dependent competition for food. Although prey availability strongly influences sea otter carrying capacity, both intrinsic and extrinsic factors influence population equilibrium density, resulting in spatiotemporal variations in foraging behavior.
Translocations maintain genetic diversity and increase connectivity in sea otters, Enhydra lutris
Sea otters, Enhydra lutris , were once abundant along the nearshore areas of the North Pacific. The international maritime fur trade that ended in 1911 left 13 small remnant populations with low genetic diversity. Subsequent translocations into previously occupied habitat resulted in several reintroduced populations along the coast of North America. We sampled sea otters between 2008 and 2011 throughout much of their current range and used 19 nuclear microsatellite markers to evaluate genetic diversity, population structure, and connectivity between remnant and reintroduced populations. Average genetic diversity within populations was similar: observed heterozygosity 0.55 and 0.53, expected heterozygosity 0.56 and 0.52, unbiased expected heterozygosity 0.57 and 0.52, for reintroduced and remnant populations, respectively. Sea otter population structure was greatest between the Northern and Southern sea otters with further structuring in Northern sea otters into Western, Central, and Southeast populations (including the reintroduced populations). Migrant analyses suggest the successful reintroductions and growth of remnant groups have enhanced connectivity and gene flow between populations throughout many of the sampled Northern populations. We recommend that future management actions for the Southern sea otter focus on future reintroductions to fill the gap between the California and Washington populations ultimately restoring gene flow to the isolated California population.
Ecosystem response persists after a prolonged marine heatwave
Some of the longest and most comprehensive marine ecosystem monitoring programs were established in the Gulf of Alaska following the environmental disaster of the Exxon Valdez oil spill over 30 years ago. These monitoring programs have been successful in assessing recovery from oil spill impacts, and their continuation decades later has now provided an unparalleled assessment of ecosystem responses to another newly emerging global threat, marine heatwaves. The 2014–2016 northeast Pacific marine heatwave (PMH) in the Gulf of Alaska was the longest lasting heatwave globally over the past decade, with some cooling, but also continued warm conditions through 2019. Our analysis of 187 time series from primary production to commercial fisheries and nearshore intertidal to offshore oceanic domains demonstrate abrupt changes across trophic levels, with many responses persisting up to at least 5 years after the onset of the heatwave. Furthermore, our suite of metrics showed novel community-level groupings relative to at least a decade prior to the heatwave. Given anticipated increases in marine heatwaves under current climate projections, it remains uncertain when or if the Gulf of Alaska ecosystem will return to a pre-PMH state.
Spatial epidemiological patterns suggest mechanisms of land-sea transmission for Sarcocystis neurona in a coastal marine mammal
Sarcocystis neurona was recognised as an important cause of mortality in southern sea otters (Enhydra lutris nereis) after an outbreak in April 2004 and has since been detected in many marine mammal species in the Northeast Pacific Ocean. Risk of S. neurona exposure in sea otters is associated with consumption of clams and soft-sediment prey and is temporally associated with runoff events. We examined the spatial distribution of S. neurona exposure risk based on serum antibody testing and assessed risk factors for exposure in animals from California, Washington, British Columbia and Alaska. Significant spatial clustering of seropositive animals was observed in California and Washington, compared with British Columbia and Alaska. Adult males were at greatest risk for exposure to S. neurona, and there were strong associations with terrestrial features (wetlands, cropland, high human housing-unit density). In California, habitats containing soft sediment exhibited greater risk than hard substrate or kelp beds. Consuming a diet rich in clams was also associated with increased exposure risk. These findings suggest a transmission pathway analogous to that described for Toxoplasma gondii, with infectious stages traveling in freshwater runoff and being concentrated in particular locations by marine habitat features, ocean physical processes, and invertebrate bioconcentration.
Trends and carrying capacity of sea otters in Southeast Alaska
Sea otter populations in Southeast Alaska (SEAK) have increased dramatically from fewer than 500 translocated animals in the late 1960s. The recovery of sea otters to ecosystems from which they had been absent has affected coastal food webs, including commercially important fisheries, and thus information on expected growth and equilibrium abundances can help inform resource management. We compile available survey data for SEAK and fit a Bayesian state-space model to estimate past trends and current abundance. Our model improves upon previous analyses by partitioning and quantifying sources of estimation error, accounting for over-dispersion of aerial count data, and providing realistic measurements of uncertainty around point estimates of abundance at multiple spatial scales. We also provide the first estimates of carrying capacity (K) for SEAK, at both regional and sub-regional scales, and analyze growth rates, current population status and expected future trends. At the regional scale, the population increased from 13,221 otters in 2003 (95% credible interval 9,990 – 16,828) to 25,584 otters in 2011 (CI95 18,739 – 33,163). The average annual growth rate in southern SEAK (7.8%) was higher than northern SEAK (2.7%); however, growth varied at the sub-regional scale and there was a negative relationship between growth rates and the number of years sea otters were present in an area. Local populations vary in terms of current densities and expected future growth: the mean estimated density at K was 4.2 (1.58) sea otters per km2 of habitat (defined as the sub-tidal benthos between 0-40m depth) and current densities correspond on average to 50% of projected equilibrium values (range = 1% to 97%) with the earliest-colonized sub-regions tending to be closer to K. Assuming a similar range of equilibrium densities for currently un-occupied habitats in SEAK, the projected value of K for all of SEAK is 74,650 sea otters (CI95 =36,778–136,506). Future analyses can improve upon the precision of K estimates by employing more frequent surveys at index sites and incorporating environmental covariates into the process model to generate habitat-specific estimates of equilibrium density.
Future directions in sea otter research and management
The conservation and management of sea otters has benefited from a dedicated research effort over the past 60 years enabling this species to recover from a few thousand in the early 20th century to about 150,000 today. Continued research to allow full, pre-exploitation recovery and restoration of nearshore ecosystems should focus on at least seven key challenges: 1) Defining sea otter populations at smaller spatial scales that reflect this species’ life history and dispersal patterns; 2) Understanding factors that regulate sea otter population density with a focus on index sites that are representative of the variety of littoral habitats occupied by sea otters around the North Pacific Rim; 3) Quantifying the effects of sea otters on the littoral community with a focus on how food availability limits population and ecosystem recovery and on predicting the effect of sea otter reoccupation on commercially valuable invertebrates; 4) Making sea otter monitoring programs comparable across geo-political boundaries through international collaboration to optimize survey efforts both spatially and temporally and to determine the cause of changes in sea otter demographics; 5) Evaluating the conservation benefits of sea otter reintroductions into historical habitat; 6) Assessing the socioeconomic costs and benefits of sea otter range expansion to anticipate and mitigate conflicts; 7) Recognizing in conservation and management plans that sea otters can be significantly affected by higher level predators in some circumstances. Many of these challenges will require new tools including next generation geolocation tag technology that will allow assessments of long-range movements, dispersal and gene flow in various populations.