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Brian B. Hatfield

Publications and source records attributed to Brian B. Hatfield.

16 recordsLinked to original sources

Dynamic and context-dependent keystone species effects in kelp forests

Sea otters are an iconic keystone predator that can maintain kelp forests by preying on grazing invertebrates such as sea urchins. However, the effects of sea otters on kelp forests vary over their geographic range. Here, we analyze two 30-y datasets on kelp forest communities during the reintroduction of sea otters along the west coast of Vancouver Island, BC, Canada, and around San Nicolas Island, CA. We developed a community model to estimate species interactions as dynamic rates, varying with community state. We find evidence of a classic trophic cascade off Vancouver Island; the arrival of otters quickly led to depletion of urchins and recovery of kelp. However, this cascade was muted around San Nicolas Island, with otters, urchins, and kelp all coexisting at intermediate densities for multiple years. Our models show that this difference came from a pulse of strong otter impacts on urchins following recolonization off Vancouver Island, but not off San Nicolas Island. The mean effects of otters on urchins and urchins on kelp were not stronger in the north, indicating that interaction dynamics and not average interaction strength are key to explaining differences in community trajectories. We also find stronger multistep interaction chains in the south, arising from competitive interactions that indirectly buffered otter effects. These findings shed light on long-standing hypotheses about how interspecific interactions can alter the function of keystone species across community contexts. More broadly, we show how community change can be more accurately predicted by considering dynamic interaction strengths.

Proceedings of the National Academy of Sciences

Southern (California) sea otter population status and trends at San Nicolas Island, 2020–2023

The population of southern sea otters ( Enhydra lutris nereis ) at San Nicolas Island, California, has been monitored annually since the translocation of 140 southern sea otters to the island was completed in 1990. Monitoring efforts have varied in frequency and type across years. In 2017, the U.S. Navy and the U.S. Fish and Wildlife Service initiated a southern sea otter monitoring and research plan to determine the effects of military readiness activities on the growth or decline of the southern sea otter population at San Nicolas Island. The southern sea otter is the only subspecies of sea otter in California (hereafter, “sea otter"). The monitoring program, at its basic level, includes seasonal surveys of population abundance, distribution, and foraging activity. From 2020 to 2023, we measured a 10-percent per annum increase in population abundance (95-percent confidence interval =0–20 percent), with 146 total individuals as of April 2023. Coinciding with the recent population growth, the sea otter distribution, which previously tended to concentrate on the island’s west end during 2003–2006 before shifting toward more use in the north and south sides during 2017–2019, appears to have shifted again during 2020–2023 to concentrate at the island’s east end. Forage data were collected between February 2020 and April 2023. There was a total of 773 forage dives in 60 forage bouts, with most of the identified prey on successful dives (n=401) recorded as sea urchins (66 percent), followed by bivalves (15 percent), snails (12 percent), and crabs (5.2 percent). Two lobsters and three abalone also were identified among the sea otter prey. Estimates of energy intake rates averaged 14.0 kilocalories per minute (95-percent confidence interval =10.8–17.2 kilocalories per minute). Monitoring data from the past two decades indicate that sea otters at San Nicolas Island have maintained a steady pattern of energy intake and population growth characteristic of a robust population, including a sixfold growth between 2000 and 2023. There was no conclusive evidence of density-dependent effects based on these patterns; however, estimates of energy intake rates for 2020–2023 were slightly lower than previous estimates from 2017 to 2019. Additionally, subtidal monitoring results at four sites around San Nicolas Island indicated that counts of purple sea urchins ( Strongylocentrotus purpuratus ) have increased between 2003 and 2023, whereas sea otter foraging surveys completed during the same period revealed that some sea otters have shifted toward higher consumption of purple sea urchins and bivalves compared to red sea urchins ( S. fransicanus ), which generally are the preferred larger prey of sea otters. These results contribute to the understanding of population dynamics and to the conservation and planning of future monitoring and research of sea otters at San Nicolas Island.

California

An integrated population model for southern sea otters

Southern sea otters ( Enhydra lutris nereis ) have recovered slowly from their near extinction a century ago, and their continued recovery has been challenged by multiple natural and anthropogenic factors. Development of an integrated population model (IPM) for southern sea otters has been identified as a management priority, to help in evaluating the relative impacts of known threats and guide best management options for species recovery. An IPM represents an analytical modeling framework where various types of data relevant to animal health, population trends, and survival can be evaluated collectively to project future population dynamics under different resource management scenarios. Here, we describe the development of a spatially explicit IPM for southern sea otters that is fit by using Bayesian methods to multiple datasets including a time series of range-wide survey counts, estimated survival rates of tagged animals from telemetry-based population studies, and cause-of-death data from comprehensive necropsies of beach-cast carcasses. The core of the model is a stage-structured matrix, in which survival rates for a given life history stage, year, and location are computed as the outcome of multiple ‘competing risks,’ or hazards, allowing for spatiotemporal variation in each hazard, density-dependence, and stochasticity. The parameterized IPM was used to (1) examine how age and sex-specific hazards vary over space and time, (2) gain insights into density-dependent variation in specific hazards, (3) assess population-level effects of known mortality hazards in the past and in future projections, and (4) evaluate the relative benefits of various potential management actions to address these hazards. Our results indicated that different types of hazards have variable impacts at different life history stages of sea otters; for example, shark-bite mortality had a strong impact on mortality of subadult females but relatively low impacts on aged adult female survival, whereas End Lactation Syndrome showed just the opposite age-based pattern. There also was spatial and temporal variation in exposure to different hazards; for example, shark-bite mortality generally was highest at the north and south ends of the sea otter range, End Lactation Syndrome and cardiac disease were highest in the center part of the range, and harmful algal bloom intoxication and protozoal infection mortalities were highest around Morro Bay. The relative impacts of hazards depended on population density; for example, shark-bite mortality had the greatest effect on male survival when population abundance was low, but as densities increased the impacts of cardiac disease (for aged adults) and acanthocephalan peritonitis (for subadults) exceeded the effects of shark-bite mortality. Sensitivity analyses showed that modifying certain hazard rates can have substantial impacts on future population growth; for example, if the shark-bite hazard rate were to decrease by 20 percent, projected abundance after 50 years is predicted to be 18-percent higher, on average, than under baseline conditions. We used the IPM to evaluate the possible impacts of a potential management action: the reintroduction of sea otters to currently unoccupied parts of their historical range. We found that there were large increases in expected growth potential associated with reintroduction programs to various locations to the north and south of the currently occupied range, although a reintroduction to San Francisco Bay was projected to have the greatest potential impacts on future population growth. The IPM for southern sea otters presented here provides resource managers with a useful tool for evaluating the impacts of specific hazards, forecasting future population dynamics and range expansion, and evaluating alternative management scenarios.

California

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.

Alaska

Habitat features predict carrying capacity of a recovering marine carnivore

The recovery of large carnivore species from over‐exploitation can have socioecological effects; thus, reliable estimates of potential abundance and distribution represent a valuable tool for developing management objectives and recovery criteria. For sea otters ( Enhydra lutris ), as with many apex predators, equilibrium abundance is not constant across space but rather varies as a function of local habitat quality and resource dynamics, thereby complicating the extrapolation of carrying capacity ( K ) from one location to another. To overcome this challenge, we developed a state‐space model of density‐dependent population dynamics in southern sea otters ( E. l. nereis ), in which K is estimated as a continuously varying function of a suite of physical, biotic, and oceanographic variables, all described at fine spatial scales. We used a theta‐logistic process model that included environmental stochasticity and allowed for density‐independent mortality associated with shark bites. We used Bayesian methods to fit the model to time series of survey data, augmented by auxiliary data on cause of death in stranded otters. Our model results showed that the expected density at K for a given area can be predicted based on local bathymetry (depth and distance from shore), benthic substrate composition (rocky vs. soft sediments), presence of kelp canopy, net primary productivity, and whether or not the area is inside an estuary. In addition to density‐dependent reductions in growth, increased levels of shark‐bite mortality over the last decade have also acted to limit population expansion. We used the functional relationships between habitat variables and equilibrium density to project estimated values of K for the entire historical range of southern sea otters in California, USA, accounting for spatial variation in habitat quality. Our results suggest that California could eventually support 17,226 otters (95% CrI = 9,739–30,087). We also used the fitted model to compute candidate values of optimal sustainable population abundance (OSP) for all of California and for regions within California. We employed a simulation‐based approach to determine the abundance associated with the maximum net productivity level (MNPL) and propose that the upper quartile of the distribution of MNPL estimates (accounting for parameter uncertainty) represents an appropriate threshold value for OSP. Based on this analysis, we suggest a candidate value for OSP (for all of California) of 10,236, which represents 59.4% of projected K .

California

Southern (California) sea otter population status and trends at San Nicolas Island, 2017–2020

The southern sea otter ( Enhydra lutris nereis ) population at San Nicolas Island, California, has been monitored annually since the translocation of 140 sea otters to the island was completed in 1990. Monitoring efforts have varied in frequency and type across years. In 2017, the U.S. Navy and the U.S. Fish and Wildlife Service initiated a sea otter monitoring and research plan to determine the effects of military readiness activities on the growth or decline of the southern sea otter population at San Nicolas Island. The monitoring program, at its basic level, includes quarterly seasonal surveys of population abundance, distribution, and foraging activity. From 2017 to 2020, we measured a 22-percent per annum increase in population abundance (95-percent confidence interval =11–34 percent) with 114 total individuals as of February 2020. Coinciding with recent population growth, the sea otter distribution, which previously tended to concentrate on the west side, appears to have shifted toward an expansion of use in the north and especially greater seasonal use in the north and south during winter and spring. Foraging data were collected on a total of 2,675 foraging dives in 167 foraging bouts, and the majority of identified prey on successful dives (n=1,335) were sea urchins (940) followed by snails (240) and crabs (78). Small numbers of lobsters (26), octopus (16), and abalone (5) also were identified. Estimates of energy intake rates averaged 17.3 kilocalories per minute (95-percent confidence interval =15.6–19.0 kilocalories per minute) and suggest possible variations across years and seasons, but confidence intervals based on specific years of data were relatively wide. In addition to abundance, trends, distribution, and forage energy intake across seasons and years, these replicated surveys provide information on the precision of data achieved by quarterly survey effort. We used precision estimates and conducted simulation analyses to assess the power of detecting 10-percent or greater decreases in population growth rates and how this power is likely to change with years of observation, survey effort, and the size of decrease. These results can be useful to the planning of future monitoring and research of sea otters at San Nicolas Island.

California

California sea otter ( Enhydra lutris nereis ) census results, spring 2019

The 2019 census of southern sea otters ( Enhydra lutris nereis ), also known as California sea otters, was conducted from early May to early July along the mainland coast of central California and in April at San Nicolas Island in southern California. The range-wide index, defined as the 3-year average of combined counts from the mainland range and San Nicolas Island, was 2,962, a decrease of 166 sea otters from the index for the previous year. The 5-year average trend in the range-wide index, including both the mainland range and San Nicolas Island populations, was flat at 0.12 percent growth per year. The northern range boundary was not calculated because of limitations in survey resources required to survey areas north of the range from the previous year (2018); however, the southern range boundary expanded slightly by 0.5 kilometer, where a negligible trend in sea otter counts (0.55 percent growth per year) corresponded to the lack of meaningful population range expansion at the southern periphery.

California

Drift and beaching patterns of sea otter carcasses and car tire dummies

Enumerating and examining marine animal carcasses is important for quantifying mortality rates and determining causes of mortality. Drifter experiments are one tool for estimating at‐sea mortality and determining factors affecting carcass drift, but they require validation to confirm drifters accurately replicate the drift characteristics of the species of interest. The goal of this study was to determine whether dummies constructed from car tires were appropriate substitutes for sea otter ( Enhydra lutris ) carcasses. We released 33 sets of targets (carcasses and dummies) in a one‐to‐one ratio on 15 randomly chosen dates between January 1995 and December 1996. They were telemetrically tracked until they beached or were no longer detected. Beaching rates were similar between carcasses (69.7%) and dummies (66.7%). Our results indicated that there was no statistical difference in the drifting pattern, as measured by distance traveled and location, between carcasses and dummies, and that cumulative wind speed, days since release, and release month were predictors of drift patterns. We concluded that dummies constructed from car tires do imitate sea otter carcasses and could be used to estimate at‐sea mortality of sea otters, or, if released during or after an oil spill, could be used to direct search efforts for carcasses.

California

California sea otter (Enhydra lutris nereis) census results, spring 2018

The 2018 census of southern sea otters ( Enhydra lutris nereis ) was conducted from late April to mid-May along the mainland coast of central California and in April at San Nicolas Island in southern California. The 3-year average of combined counts from the mainland range and San Nicolas Island was 3,128, a decrease of 58 sea otters from the previous year. The 5-year average trend in abundance, including both the mainland range and San Nicolas Island populations, remains positive at 1.3 percent per year. Continuing lack of growth in the range peripheries likely explains the cessation of range expansion.

California

California sea otter (Enhydra lutris nereis) census results, Spring 2017

The 2017 census of southern sea otters ( Enhydra lutris nereis ) was conducted between late April and early July along the mainland coast of central California and in April at San Nicolas Island in southern California. The 3-year average of combined counts from the mainland range and San Nicolas Island was 3,186, down by 86 sea otters from the previous year. This is the second year that the official index has exceeded 3,090, the Endangered Species Act delisting threshold identified in the U.S. Fish and Wildlife Service’s Southern Sea Otter Recovery Plan (the threshold would need to be exceeded for 3 consecutive years before delisting consideration). The 5-year average trend in abundance, including both the mainland range and San Nicolas Island populations, remains positive at 2.3 percent per year. Continuing lack of growth in the range peripheries likely explains the cessation of range expansion.

California

California sea otter ( Enhydra lutris nereis ) census results, Spring 2016

The 2016 census of southern sea otters Enhydra lutris nereis was conducted in May along the mainland coast of central California and in April at San Nicolas Island in southern California. The 3-year average of combined counts from the mainland range and San Nicolas Island was 3,272. This is the first year that the official index has exceeded 3,090, the Endangered Species Act delisting threshold suggested by the U.S. Fish and Wildlife Service (the threshold would need to be exceeded for 3 consecutive years before delisting consideration). The 5-year average trend in abundance, including both the mainland range and San Nicolas Island populations, is positive at 3.2 percent per year; however, regional trends vary, with localized declines at the southern and northern peripheries of the mainland range. The lack of population growth in the range peripheries over recent years likely explains the cessation of range expansion, with the range limits remaining almost unchanged from the previous 5 years.

California

Dramatic increase in sea otter mortality from white sharks in California

Although southern sea otters (Enhydra lutris nereis) are not considered prey for white sharks (Carcharodon carcharias), sharks do nonetheless bite sea otters. We analyzed spatial and temporal trends in shark bites on sea otters in California, assessing the frequency of shark bite wounds in 1,870 carcasses collected since 1985. The proportion of stranded sea otters having shark bites has increased sharply since 2003, and white shark bites now account for >50% of recovered carcasses. The trend was most pronounced in the southern part of the range, from Estero Bay to Point Conception, where shark bite frequency has increased eightfold. Seasonal trends were also evident: most shark-bitten carcasses are recovered in late summer and fall; however, the period of elevated shark bite frequency has lengthened. The causes of these trends are unclear, but possible contributing factors include increased white shark abundance and/or changes in white shark behavior and distribution. In particular, the spatiotemporal patterns of shark-bitten sea otters match increases in pinniped populations, and the increased availability of marine mammal prey for white sharks may have led to more sharks spending more time in nearshore waters utilized by both sea otters and pinnipeds.

California

Sea otter mortality in fish and shellfish traps: Estimating potential impacts and exploring possible solutions

Sea otters Enhydra lutris can be bycaught and drowned in fishing pots and traps, which may pose a threat to the welfare of otter populations. We explored this potential problem and its solutions using a wide variety of analyses. We exposed live California (USA) sea otters to finfish traps, lobster traps, and mock Dungeness crab traps in captive trials and found that the animals attempted to enter the circular and rectangular fyke openings, with some becoming entrapped. Using both live and dead sea otters, we found that a 3 × 9 inch (7.6 × 22.9 cm) fyke opening (1 inch narrower than the 4 × 9 inch [10.2 × 22.9 cm] openings currently used in California’s commercial Dungeness crab fishery) would exclude most free-living (i.e. weaned from their mothers) otters while permitting the undiminished capture of crabs. Observer programs do not currently exist in California for these fisheries, so we calculated the effort required by an observer program to document sea otter bycatch over a range of hypothetical levels and evaluated the impact of those mortality rates on population growth. These analyses demonstrate that significant mortality from bycatch might easily go undetected, even with seemingly high levels of observer effort. As sea otters reoccupy portions of their former habitat in California, co-occurrence with finfish and shellfish traps with relatively large fyke openings will increase.

Endangered Species Research

Interactions between northern elephant seals and vehicles near Point Piedras Blancas, California

Northern elephant seals, Mirounga angustirostris , were nearly extirpated in the late 19th and early 20th centuries. However, they have recovered to the point where they are now locally abundant (Stewart e t al. 1994). Stimulated by the report of human/northern elephant seal interactions in Baja California Sur (Webster and Baird 1998), we report here on an unusual terrestrial example of the increasing conflict between the growing populations of these two species in central California.

California

Incorporating diverse data and realistic complexity into demographic estimation procedures for sea otters

Reliable information on historical and current population dynamics is central to understanding patterns of growth and decline in animal populations. We developed a maximum likelihood-based analysis to estimate spatial and temporal trends in age/sex-specific survival rates for the threatened southern sea otter (Enhydra lutris nereis), using annual population censuses and the age structure of salvaged carcass collections. We evaluated a wide range of possible spatial and temporal effects and used model averaging to incorporate model uncertainty into the resulting estimates of key vital rates and their variances. We compared these results to current demographic parameters estimated in a telemetry-based study conducted between 2001 and 2004. These results show that survival has decreased substantially from the early 1990s to the present and is generally lowest in the north-central portion of the population's range. The greatest temporal decrease in survival was for adult females, and variation in the survival of this age/sex class is primarily responsible for regulating population growth and driving population trends. Our results can be used to focus future research on southern sea otters by highlighting the life history stages and mortality factors most relevant to conservation. More broadly, we have illustrated how the powerful and relatively straightforward tools of information-theoretic-based model fitting can be used to sort through and parameterize quite complex demographic modeling frameworks. ?? 2006 by the Ecological Society of America.

Ecological Applications

Status of translocated sea otters at San Nicolas Island, California

In the 1970s about 1,650 southern sea otters (Enhydra lutris nereis) were restricted to the central California coast (Riedman and Estes, 1990), and a high volume of oil was being shipped through the region. Because of the vulnerability of sea otters to contamination from oil (Costa and Kooyman, 1982; Williams and Davis, 1995) that would likely spread wide- ly along the shore after a large spill (Van- Blaricom and Jameson, 1982), the subspecies was listed as threatened in 1977 under the United States Endangered Species Act.

Southwestern Naturalist