Geology topics
Kimberly A. Kloecker
Publications and source records attributed to Kimberly A. Kloecker.
Quantifying sea otter abundance, distribution, habitat use, and foraging intake in Cook Inlet, Alaska
Following near extirpation from the fur trade, sea otters ( Enhydra lutris ) have returned to occupy lower Cook Inlet since the 1950s, or earlier, with numbers increasing to ~11,000 and ~9,000 on the west and east side, respectively, by 2017. Northward range expansion on the west side has been negligible for decades with few animals found north of Kamishak Bay, while northward expansion on the east side has been more pronounced in recent decades. The reasons for these contrasting distribution patterns are not certain. Possible explanations for lack of expansion on the west side included 1) poor sea otter habitat north of Kamishak Bay; 2) adequate habitat north of Kamishak Bay but no incentive for sea otters to move north because of abundant food in Kamishak Bay, and/or sea otters discouraged from moving north of Kamishak Bay; 3) seasonal ice formation; or 4) seasonal presence of killer whales. This project was designed to document current sea otter abundance and distribution in lower Cook Inlet, including seasonal variation, and evaluate drivers of habitat use and foraging conditions, including how these overlap with Bureau of Ocean Energy Management (BOEM) Lease Sale blocks. We found that the probability of sea otter presence in lower Cook Inlet was significantly related to depth and distance from shore with the highest probabilities of sea otter presence in areas ~7–8 kilometers (km) from shore in water ~20–30 meters (m) deep. Historical survey data suggest sea otter distribution has not changed dramatically since 2002, and while we detected significant seasonal changes including due to presence of heavy sea ice, the seasonal changes were modest and primarily related to locations with consistent winter ice formation on the west side of lower Cook Inlet. Overall, throughout the year, sea otters appeared to be utilizing most of the habitat within the 40-m depth contour on the east side of lower Cook Inlet, including Kachemak Bay. Sea otters on the west side reside largely within Kamishak Bay. Because of the shallow bathymetry of lower Cook Inlet, sea otters consistently occurred farther offshore than in many other areas of Alaska. The presence of sea otters, including females with pups, in these offshore waters indicates that sea otters can forage productively in these open water areas. Importantly, we documented that sea otters, including females with pups, occurred in most of the eastern BOEM lease blocks including the original Lease Sale 244 blocks 7064, 7114, 6162, 6310, 6360, 6410, 6458 and 6457. The relinquishments for these 7 leases were effective September 17, 2024: OCS-Y-02434 (block # 7064), OCS-Y-02435 (7114), OCS-Y-02436 (6162), OCS-Y-02438 (6357), OCS-Y-02442 (6407), OCS-Y02446 (6457), OCS-Y-02447 (6458) https://www.boem.gov/sites/default/files/documents/environment/Map%20of%20Active%20Leases%20 Cook%20Inlet%20OCS_0.pdf Regarding differential northward expansion on the east and west side of lower Cook Inlet, we found that wind and water circulation patterns make winter sea ice more prominent on the west side, but this had minimal effects on sea otter distribution. In addition, although there have been observations of killer whale ( Orcinus orca ) predation on sea otters in Cook Inlet, we did not see behavioral or distributional evidence that it was prevalent enough to have strong effects on sea otter habitat use. Benthic surveys using a remotely operated vehicle (ROV) indicated that epibenthic substrate and biological community heterogeneity differed between areas with and without otters, suggesting that these factors may explain the current distribution of otters within lower Cook Inlet. However, ROV surveys cannot assess infaunal prey abundance, making assessments of the role of prey availability difficult. The existence of a healthy Pacific razor clam ( Siliqua patula ) fishery along western lower Cook Inlet along the Lake Clark coast north of Kamishak Bay suggests there may still be a resource base for eventual expansion of sea otters into this area. Our shore-based forage observations indicated that sea otters exist near carrying capacity densities relative to nearshore prey resources. However, our distribution model suggests the bulk of the population lives offshore beyond our ability to observe their feeding activity. The fact that sea otters are a consistent presence in offshore areas suggests that prey resources in these areas were relatively abundant in comparison to nearshore prey resources, making foraging in offshore areas, with water depths of 0–40 m, energetically profitable. Prey types in these offshore areas of lower Cook Inlet likely included epifauna such as crabs and large urchins, in addition to infaunal clams based on the soft substrate habitat types that characterize lower Cook Inlet. Collectively, our results suggest that sea otters occupy most areas of lower Cook Inlet with appropriate benthic habitat types and prey resources, which includes areas within BOEM Lease Sale blocks. With the exception of the Lake Clark coast, sea otters may be approaching a food- and habitat-limited distribution and carrying capacity, suggesting that the current status of sea otters in lower Cook Inlet is likely to remain similar in the absence of significant changes to prey, habitat, predation, or anthropogenic disturbance.
The dynamics of sea otter prey selection under population growth and expansion
Sea otters ( Enhydra lutris ) were extirpated from much of their range in the North Pacific by the early 1900s but have made a remarkable recovery in Southeast Alaska. Sea otter populations have been particularly successful in Glacier Bay, Alaska, a protected tidewater glacier fjord with a diverse and productive nearshore habitat. Collection of sea otter foraging observations in Glacier Bay began in 1993, along with high-resolution aerial surveys that provide estimates of sea otter abundance and distribution. We integrated these two data sources to investigate how sea otter diet changed in space and time as sea otters established and spread across Glacier Bay. Specifically, we developed a multilevel Bayesian model to capture how sea otter diet at a location (the number, type, and size of prey collected) changed as a function of local cumulative otter abundance and the year in which the location was first occupied. This framework enabled us to estimate the sequence of sea otter prey selection and switching as prey populations responded to sea otter foraging pressure. We found that local sea otter diet changed substantially as the population established, shifting away from large urchins, crabs, and clams to Modiolus mussels and small urchins, and lastly to small clams and Mytilus mussels. We also found that sea otter diet at newly occupied sites changed as otters spread over the main channel and into the arms of Glacier Bay. Further, by 2019, sea otters across the bay were primarily foraging on small prey, regardless of the local occupancy history. The absence of a spatial gradient in the size of prey captured late in the study suggests that feedbacks between the top-down effects of sea otter foraging, sea otter dispersal processes, and local variation in habitat productivity may have homogenized the size structure of available prey across Glacier Bay.
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.
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.
Nearshore ecosystems in the Gulf of Alaska
Nearshore ecosystem monitoring in western Prince William Sound, Kenai Fjords National Park, Kachemak Bay, and Katmai National Park and Preserve has been conducted as a single Nearshore Component of the Gulf Watch Alaska program over the past five years (2017-2021). This program builds on the previous five years and continues, in many cases, decades of preceding research and monitoring. During the 2017-2021 period of Gulf Watch Alaska, we successfully collected data on more than 200 nearshore metrics across all regions. These metrics were explicitly selected because of their value as trophically-connected features that offer insights into causes of changes through bottom-up and top-down forces within the nearshore food web. During these past 5 years, this study design has been applied to documenting and understanding several perturbations, including the Pacific Marine Heatwave and loss of sea stars due to sea star wasting. Continued monitoring will allow for a better understanding of change in nearshore ecosystems across the Gulf of Alaska and the status of spill injured resources, including greater perspective on oil spill effects and recovery relative to other types of perturbations. This information will be critical for anticipating and responding to ongoing and future changes in the region, as well as providing data for understanding global-scale variation in marine environments.
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.
Divergent gene expression profiles in Alaskan sea otters: An indicator of chronic domoic acid exposure?
An opportunistic investigation into ecosystem instability in Kachemak Bay (KBay), Alaska, has led us to investigate exposure to toxic algae in sea otters. We used gene expression to explore the physiological health of sea otters sampled in KBay in May 2019. We found altered levels of gene transcripts in comparison with reference sea otters from clinically normal, oil-exposed, and nutritionally challenged populations sampled over the past decade. KBay sea otters were markedly divergent from the other groups for five genes, which indicated the involvement of neurological, cardiac, immune, and detoxification systems. Further, analyses of urine and fecal samples detected domoic acid in the KBay sea otters. In combination, these results may point to chronic, low-level exposure to an algal toxin, such as domoic acid. With a warming climate, the frequency and severity of harmful algal blooms in marine environments is anticipated to increase, and novel molecular technologies to detect sublethal or chronic exposure to algal toxins will help provide an early warning of threats to the stability of populations and ecosystems.
Genetic variation in sea otters (Enhydra lutris) from the North Pacific with relevance to the threatened Southwest Alaska Distinct Population Segment
For the sea otter ( Enhydra lutris ), genetic population structure is an area of research that has not received significant attention, especially in Southwest Alaska where that distinct population segment has been listed as threatened since 2005 pursuant to the U.S. Endangered Species Act. In this study, 501 samples from 14 locations from Prince William Sound, Alaska to the Commander Islands in Russia were analyzed for variation at 13 microsatellite loci. Our results indicate a high degree of genetic divergence among the 14 locations ( F ST = 0.120) with gene flow conforming to the isolation by distance (IBD) model ( r 2 = 0.491, p < .05). The 14 sampling locations formed six geographic associations in clustering and ordination analyses that likely correspond to remnant population lineages: (1) Southcentral Alaska, (2) Kodiak and North Alaska Peninsula, (3) South Alaska Peninsula and Bristol Bay, (4) Eastern Aleutian, (5) Western Aleutian, and (6) the Commander Islands. Except for South Alaska Peninsula and Bristol Bay, these clusters closely agree with previously defined stock and management unit boundaries. Our results reveal significant genetic population structure and are generally congruent with current management strategies for the threatened Southwest Alaska distinct population segment.
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.
Timelines and mechanisms of wildlife population recovery following the Exxon Valdez oil spill
Research and monitoring activities over the 28 years since the T/V Exxon Valdez ran aground and spilled oil into Prince William Sound, Alaska have led to an improved understanding of how wildlife populations were damaged, as well as the mechanisms and timelines of recovery. A key finding was that for some species, such as harlequin ducks and sea otters, chronic oil spill effects persisted for at least two decades and were a larger influence on population dynamics over the long term than acute effects of the spill. These data also offer insights into population variation resulting from factors other than the oil spill. For example, while many seabirds experienced direct and indirect effects of the spill, population trajectories of some piscivorous birds, including pigeon guillemots and marbled murrelets, were linked to long-term environmental changes independent of spill effects. Another species, killer whales, suffered population declines due to acute spill effects that have not been resolved despite lack of chronic direct effects, representing a novel pathway of long-term injury. The observed variation in mechanisms and timelines of recovery is linked to species specific life history and natural history traits, and thus may be useful for predicting population recovery for other species following other spills.
Pacific blue mussel (Mytilus trossulus) abundance in the Gulf of Alaska: Synthesis of Gulf Watch data (2006-2013) and a consideration of major recruitment events (1989-2013)
Pacific blue mussels ( Mytilus trossulus ) are abundant and wide-spread primary consumers in the intertidal zone throughout the Gulf of Alaska (GOA). As a component of the Gulf Watch Alaska monitoring program, they represent a key member of intertidal communities and an important prey resource to a number of nearshore vertebrate predators. Our goal is to understand variation in abundance of M. trossulus over large temporal and spatial scales and over a variety of habitats in the northern GOA to determine the bottom-up factors that influence recruitment and the top-down forces that control total biomass. This information is needed to predict consequences of variation due to incremental climate change, periodic regime shifts, and catastrophic change caused by oil spills or natural events such as severe winters.
Variability within nearshore ecosystems of the Gulf of Alaska
Nearshore marine habitats, which represent the interface among air, land and sea, form a critical component of the Gulf of Alaska (GOA) ecosystem. As an interface, the nearshore facilitates transfer of water, nutrients and biota between terrestrial and oceanic systems, creating zones of high productivity. The nearshore provides a variety of ecosystem services, including (1) nursery grounds for a wide variety of marine invertebrates and fishes (e.g., crabs, salmon, and herring), (2) nesting and pupping habitats for many pelagic marine predators (e.g., sea bird nesting colonies and pinniped rookeries), (3) important feeding habitats for high trophic level pelagic predators (e.g., killer whales), (4) habitat for resident nearshore species (including sea otters, harbor seals, shorebirds, sea ducks, nearshore fishes, and marine invertebrates), many of which are important sources of commercial and subsistence harvests, and (5) recreational, commercial and subsistence opportunities for human populations (Figure 1-1). The canopy forming kelps and eel grass beds found in the nearshore provide primary production and structure to nursery habitats, and also can dissipate wave energy thus reducing coastal erosion, and serve as a carbon “sink” capable of storing substantial amounts of atmospheric CO 2 (Wilmers et al. 2012).
Exxon Valdez Oil Spill Restoration Project final report: Monitoring for evaluation of recovery and restoration of injured nearshore resources
In 2012, we completed three consecutive years of full field sampling in WPWS for EVOS Restoration Project 10100750. Nearshore monitoring was conducted in collaboration with the NPS SWAN I&M program and, beginning in 2012, as part of the EVOSTC GWA program. Data collection was done in accordance with standard operating procedures set forth to monitor marine water chemistry and quality, marine intertidal invertebrates, kelps and seagrasses, marine birds, black oystercatchers, and sea otters. Summer sampling in 2012 represented the fourth year of sampling in WPWS (an initial year of sampling was done in WPWS in 2007; EVOS Restoration Project 070750). Based on our monitoring of nearshore species in WPWS, and comparisons of data from WPWS and other areas within the Gulf of Alaska, we have no evidence of continued injury to biological resources at the spatial scales we are monitoring. A key finding is that recovery of the sea otter population is no longer constrained by exposure to lingering oil; this is consistent with related EVOSTC studies on harlequin ducks (Restoration Project 12120114-Q). We anticipate continued annual nearshore monitoring in WPWS and at KATM and KEFJ under GWA, with data summaries and analyses including all three areas to provide a larger spatial and temporal context to the understanding of processes and patterns in nearshore ecosystems of the GOA which were impacted by the EVOS of 1989.
2013 update on sea otter studies to assess recovery from the 1989 Exxon Valdez oil spill, Prince William Sound, Alaska
On March 24, 1989, the tanker vessel Exxon Valdez ran aground in Prince William Sound, Alaska, spilling an estimated 42 million liters of Prudhoe Bay crude oil. Oil spread in a southwesterly direction and was deposited on shores and waters in western Prince William Sound (WPWS). The sea otter (Enhydra lutris) was one of more than 20 nearshore species considered to have been injured by the spill. Since 1989, the U.S. Geological Survey has led a research program to evaluate effects of the spill on sea otters and assess progress toward recovery, as defined by demographic and biochemical indicators. Here, we provide an update on the status of sea otter populations in WPWS, presenting findings through 2013. To assess recovery based on demographic indicators, we used aerial surveys to estimate abundance and annual collections of sea otter carcasses to evaluate patterns in ages-at-death. To assess recovery based on biochemical indicators, we quantified transcription rates for a suite of genes selected as potential indicators of oil exposure in sea otters based on laboratory studies of a related species, the mink (Mustela vison). In our most recent assessment of sea otter recovery, which incorporated results from a subset of studies through 2009, we concluded that recovery of sea otters in WPWS was underway. This conclusion was based on increasing abundance throughout WPWS, including increasing numbers at northern Knight Island, an area that was heavily oiled in 1989 and where the local sea otter population had previously shown protracted injury and lack of recovery. However, we did not conclude that the WPWS sea otter population had fully recovered, due to indications of continuing reduced survival and exposure to lingering oil in sea otters at Knight Island, at least through 2009. Based on data available through 2013, we now conclude that the status of sea otters—at all spatial scales within WPWS—is consistent with the designation of recovery from the spill as defined by the Exxon Valdez Oil Spill Trustee Council. The support for this conclusion is based primarily on demographic data, including (1) a return to estimated pre-spill abundance of sea otters at northern Knight Island, and (2) a return to pre-spill mortality patterns. Gene transcription rates in 2012 were similar in sea otters from oiled, moderately oiled and unoiled areas, suggesting abatement of exposure effects in 2012. However, because 2012 gene transcription rates generally were low for sea otters from all areas relative to 2008, we cannot fully interpret these observations without data from a wider panel of genes. This slight uncertainty with respect to the data from the biochemical indicator is outweighed by the strength of the data for the demographic indicators. The return to pre-spill numbers and mortality patterns suggests a gradual dissipation of lingering oil over the past two decades, to the point where continuing exposure is no longer of biological significance to the WPWS sea otter population.
Synthesis of nearshore recovery following the 1989 Exxon Valdez oil spill: sea otter liver pathology and survival in Western Prince William Sound, 2001 – 2008
We examined livers and liver biopsies collected from captured sea otters in WPWS, 2001–2008, to determine whether indicators of liver health correlated with history of oil contamination from the 1989 Exxon Valdez oil spill. Sea otters captured in oiled areas had a significantly higher proportion of livers with gross pathological change, based on visual inspection at the time of capture, than those from unoiled areas. Of the 10 histopathology variables scored on liver biopsies, only two (vacuolar change and pigment) differed between animals from oiled and unoiled areas, and neither correlated with gross pathology scores. Vacuolar change indicates physiological disturbance, which is consistent with potential effects from oil exposure but also could be influenced by a number of other factors. We concluded that, as of 2008, some differences in liver health were evident between sea otters from oiled and unoiled areas; these differences were consistent with, but not specific to, effects that might be expected with sublethal exposure to lingering Exxon Valdez oil. We also quantified variation in survival of radiomarked sea otters within oiled areas of WPWS in relation to age, sex, body condition, selected blood serum chemistry variables, and histological scores indicative of liver health. Of the variables considered, only the serum enzyme aspartate aminotransferase (AST) and the ratio of serum proteins albumin and globulin (A/G) were correlated with survival, with higher levels of AST and lower levels of A/G associated with increased likelihood of mortality. High AST and low A/G both may be indicative of liver disease. Taken together, results reported here suggest that liver health of sea otters in oiled areas was slightly poorer than those from unoiled areas and, ifurther, that this may have translated to poorer survival through 2008, nearly 2 decades after the spill. More recently collected information indicated that mortality patterns and abundance had returned to pre-spill conditions between 2010 and 2013, suggesting that the effects that we detected through 2008 may have represented the end of effects related to exposure to lingering oil.