Ten years later: An update on the status of collections of endemic Gulf of Mexico fishes put at risk by the 2010 Oil Spill.

The 2010 Gulf of Mexico Deepwater Horizon was the largest oil spill in human history that occurred during a 12-week period in a region less than 100 km from the coast of Louisiana; however, after more than a decade of post-spill research, few definitives can be said to be known about the long-term impacts on the development and distribution of fishes in and around the region of the disaster. Here, we examine endemic Gulf of Mexico fish species that may have been most impacted by noting their past distributions in the region of the spill and examining data of known collecting events and observations over the last twenty years (ten years prior to the spill, ten years post-spill). Five years post-spill, it was reported that 48 of the Gulf's endemic fish species had not been collected and, with expanded methods, we now report that 29 (of the 78 endemic species) have not been reported in collections since 2010 (five of these are only known from observations post-spill). Although the good news that some previously 'missing' species have been found may be cause to celebrate, the lack of information for many species remains a cause for concern given focused sampling efforts post-spill.

Spatiotemporal patterns in the prevalence of microscopic hepatic changes in Gulf of Mexico Tilefish (Lopholatilus chamaeleonticeps) and associations with hepatic PAHs.

Following the 2010 Deepwater Horizon blowout, demersal longline surveys were conducted across the Gulf of Mexico (GoM) continental shelf to evaluate polycyclic aromatic hydrocarbon (PAH) exposure, tissue accumulation, and health indices in demersal fishes. Tilefish (Lopholatilus chamaeleonticeps), a target species due to Gulf-wide distribution with documented high exposure to PAHs, were collected in the north central GoM at repeat stations 2012 to 2015, and from the northwest GoM, Bay of Campeche, and Yucatán Shelf in 2015 and 2016. Liver samples (n = 239) were analyzed for microscopic hepatic changes (MHCs) by a board-certified veterinary pathologist. Histological analyzes identified 14 MHCs. Prevalence of MHCs was generally uniform throughout the GoM, except for low prevalence on the Yucatán Shelf. Inflammatory and vacuolar changes were most prevalent, while pre-neoplasia and neoplasia were rare. Tilefish sampled annually in the north central GoM showed increases in inflammatory MHCs and glycogen-type vacuolar change over time, while lipid-type vacuolar change decreased over time. Short-term exposure to PAHs was assessed by measuring PAH metabolites in bile (n = 100) using high performance liquid chromatography with fluorescence detection. Longer-term accumulation of PAHs in tissue was assessed by analyzing liver (n = 111) for PAHs and alkylated homologs using QuEChERS extractions and gas chromatography tandem mass spectrometry. Six MHCs including glycogen-type vacuolar change, biliary fibrosis, foci of cellular alteration, parasites, hepatocellular atrophy, and necrosis were significantly associated with hepatic PAH accumulation in Tilefish from the northern central GoM; however, no MHCs were associated with biliary PAH metabolites. Combined with previous studies of PAH exposure and health indices in north central GoM Tilefish post-Deepwater Horizon, which also identified decreases in hepatic lipid storage and Fulton's condition factor that were correlated to increasing PAH exposure, these data indicate concerning temporal trends and changes in hepatic energy storage.

Brief Oil Exposure Reduces Fitness in Wild Gulf of Mexico Mahi-Mahi (Coryphaena hippurus).

The Deepwater Horizon (DWH) disaster released 3.19 million barrels of crude oil into the Gulf of Mexico (GOM) in 2010, overlapping the habitat of pelagic fish populations. Using mahi-mahi (Coryphaena hippurus)─a highly migratory marine teleost present in the GOM during the spill─as a model species, laboratory experiments demonstrate injuries to physiology and behavior following oil exposure. However, more than a decade postspill, impacts on wild populations remain unknown. To address this gap, we exposed wild mahi-mahi to crude oil or control conditions onboard a research vessel, collected fin clip samples, and tagged them with electronic tags prior to release into the GOM. We demonstrate profound effects on survival and reproduction in the wild. In addition to significant changes in gene expression profiles and predation mortality, we documented altered acceleration and habitat use in the first 8 days oil-exposed individuals were at liberty as well as a cessation of apparent spawning activity for at least 37 days. These data reveal that even a brief and low-dose exposure to crude oil impairs fitness in wild mahi-mahi. These findings offer new perspectives on the lasting impacts of the DWH blowout and provide insight about the impacts of future deep-sea oil spills.

Hypersaline Pore Water in Gulf of Mexico Beaches Prevented Efficient Biodegradation of Deepwater Horizon Beached Oil.

The 2010 Deepwater Horizon (DWH) blowout released 3.19 million barrels (435 000 tons) of crude oil into the Gulf of Mexico. Driven by currents and wind, an estimated 22 000 tons of spilled oil were deposited onto the northeastern Gulf shorelines, adversely impacting the ecosystems and economies of the Gulf coast regions. In this work we present field work conducted at the Gulf beaches in three U.S. States during 2010-2011: Louisiana, Alabama, and Florida, to explore endogenous mechanisms that control persistence and biodegradation of the MC252-oil deposited within beach sediments as deep as 50 cm. The work involved over 1500 measurements incorporating oil chemistry, hydrocarbon-degrading microbial populations, nutrient and DO concentrations, and intrinsic beach properties. We found that intrinsic beach capillarity along with groundwater depth provides primary controls on aeration and infiltration of near-surface sediments, thereby modulating moisture and redox conditions within the oil-contaminated zone. In addition, atmosphere-ocean-groundwater interactions created hypersaline sediment environments near the beach surface at all the studied sites. The fact that the oil-contaminated sediments retained near or above 20% moisture content and were also eutrophic and aerobic suggests that the limiting factor for oil biodegradation is the hypersaline environment due to evaporation, a fact not reported in prior studies. These results highlight the importance of beach porewater hydrodynamics in generating unique hypersaline sediment environments that inhibited oil decomposition along the Gulf shorelines following DWH.

Polycyclic Aromatic Hydrocarbon-DNA Adducts in Gulf of Mexico Sperm Whale Skin Biopsies Collected in 2012.

The northern Gulf of Mexico has a long history of polycyclic aromatic hydrocarbon (PAH) contamination from anthropogenic activities, natural oil seepages, and the 2010 Deepwater Horizon explosion and oil spill. The continental shelf of the same area is a known breeding ground for sperm whales (Physeter macrocephalus). To evaluate PAH-DNA damage, a biomarker for potential cancer risk, we compared skin biopsies collected from Gulf of Mexico sperm whales in 2012 with skin biopsies collected from sperm whales in areas of the Pacific Ocean in 1999-2001. All samples were obtained by crossbow and comprised both epidermis and subcutaneous blubber. To evaluate exposure, 7 carcinogenic PAHs were analyzed in lipids extracted from Pacific Ocean sperm whale blubber, pooled by sex, and location. To evaluate PAH-DNA damage, portions of all tissue samples were formalin-fixed, paraffin-embedded, sectioned, and examined for PAH-DNA adducts by immunohistochemistry (IHC) using an antiserum elicited against benzo[a]pyrene-modified DNA, which crossreacts with several high molecular weight carcinogenic PAHs bound to DNA. The IHC showed widespread epidermal nuclear localization of PAH-DNA adducts in the Gulf of Mexico whales (n = 15) but not in the Pacific Ocean whales (n = 4). A standard semiquantitative scoring system revealed significantly higher PAH-DNA adducts in the Gulf of Mexico whales compared to the whales from the Pacific Ocean study (p = .0002).

Mapping spatial and temporal variation of seafloor organic matter Δ14C and δ13C in the Northern Gulf of Mexico following the Deepwater Horizon Oil Spill.

Following the Deepwater Horizon oil spill of 2010, large amounts of biodegraded oil (petrocarbon) sank to the seafloor. Our objectives were to 1) determine post-spill isotopic values as the sediments approached a new baseline and 2) track the recovery of affected sediments. Sediment organic carbon δ13C and Δ14C reached a post-spill baseline averaging -21.2 ± 0.9‰ (n = 129) and -220 ± 66‰ (n = 95). Spatial variations in seafloor organic carbon baseline isotopic values, 13C and 14C, were influenced by river discharge and hydrocarbon seepage, respectively. Inverse Distance Weighting of surface sediment Δ14C values away from seep sites showed a 50% decrease in the total mass of petrocarbon, from 2010 to 2014. We estimated a rate of loss of -2 × 109 g of petrocarbon-C/year, 2-11% of the degradation rates in surface slicks. Despite the observed recovery in sediments, lingering residual material in the surface sediments was evident seven years following the blowout.

A review of Gulf of Mexico coastal marsh erosion studies following the 2010 Deepwater Horizon oil spill and comparison to over 4 years of shoreline loss data from Fall 2010 to 2015.

The 2010 Deepwater Horizon (DWH) oil spill affected nearly 1105 km of coastal marsh. Long-term shoreline loss in the northern Gulf of Mexico is an important question with far-reaching ecological and human-use implications. Numerous studies have examined potential exacerbated marsh shoreline retreat after the DWH using ground-level sampling and/or aerial/satellite imagery interpretation. This paper reviews previous DWH erosion studies, discusses their limitations and sometimes conflicting results, and provides a comprehensive analysis of a larger data set. Shoreline retreat measurements from multiple studies following the DWH incident were combined for 131 herbaceous marsh sample sites for the period from Fall 2010 to Summer 2015. Significant increases in shoreline loss were found only in the period from Fall 2010 to Fall 2011 for heavily oiled shorelines relative to other periods. The evidence does not suggest widespread long-term coastal marsh erosion from the DWH.

Impacts of dibenzopyrenes on bacterial community isolated from Gulf of Mexico sediment.

The presence of polycyclic aromatic hydrocarbons (PAHs) in marine environments as a result of contamination is an environmental concern, especially in regions where oil spills such as the Deepwater Horizon have occurred. While numerous PAHs have been studied for their effects on microbes, the family of dibenzopyrenes has yet to be investigated. In this preliminary study, the impacts of these molecules on the community structure of a bacterial consortium isolated from oil-impacted Gulf of Mexico sediment were examined using high-throughput sequencing, demonstrating intriguing negative impacts on species diversity and abundance. While no measurable degradation of the dibenzopyrenes was observed after 28-day incubation, the abundance of known oil-degrading bacteria from orders such as Oceanospirillales, Caulobacterales, Sphingomonadales, and Nitrosococcales were shown to be enhanced. Of the five isomers of dibenzopyrene studied, dibenzo[a,h]pyrene supported the fewer number of microbial species suggesting the isomer was more toxic compared to the other isomers.

Evaluation of the fate of carcasses and dummies deployed in the nearshore and offshore waters of the northern Gulf of Mexico.

In response to the Deepwater Horizon oil spill, federal and state agencies conducted field studies to develop inputs for a shoreline deposition model used to estimate nearshore avian mortality resulting from the spill. A 2011 carcass drift study was designed to generate data on the likelihood that birds that died on the water would deposit along the northern Gulf of Mexico coast (rather than becoming lost at sea). In the case of the Deepwater Horizon oil spill, carcass losses at sea accounted for a significant portion of nearshore avian mortality. We evaluate the data collected during the Deepwater Horizon carcass drift study and compare the results obtained from the use of avian carcasses versus dummy carcasses (dummies) and the differences between those deployed nearshore versus further offshore. We conclude that, although the use of dummies provided valuable confirmation on the drift patterns of dead birds, dummies drifted greater distances, for longer periods of time, and were more likely to be observed beached compared to avian carcasses, with 64.6% of dummies beaching compared to 17.2% of carcasses. In response to future spills, researchers should account for these potential biases when incorporating dummy drift data into estimates of avian carcass loss. Further, none of the avian carcasses and dummies released more than 40 km from the shoreline made it to shore. In the northern Gulf of Mexico, carcasses that die on the waters farther offshore are unlikely to make it to shore to be captured in a deposition model; therefore, it may be appropriate to utilize a separate methodology to estimate offshore mortality. The applicability of these results to other spill events should be evaluated in the context of the specific spill characteristics.

Pelagic seabird density and vulnerability in the Gulf of Mexico to oiling from the Deepwater Horizon/MC-252 spill.

Using ship-based surveys, the Natural Resource Damage Assessment (NRDA) Trustees assessed the external oiling of offshore and pelagic marine birds inhabiting the northern Gulf of Mexico (Gulf) in the year following the Deepwater Horizon oil spill (DWH spill). Study objectives were to (1) collect data on pelagic seabirds that were visibly oiled, (2) collect data to estimate abundance of seabirds in offshore and pelagic waters, and 3) document the location and condition of any bird carcasses encountered. Methods employed included strip line transects and station point counts. Surveys were conducted within a study area bound by the Texas-Mexico border and the Dry Tortugas of Florida to the south, and the nearshore coastal waters of the northern Gulf of Mexico. A total of 5665 strip line transects and 386 station point-counts of variable duration were collected during the study. More than 23,000 individual seabirds comprising 45 estuarine, coastal, offshore, and pelagic species were tallied. Average daily abundance of seabirds detected varied from a low of approximately 7 birds/day in November 2010 along regions of the mid- and outer continental shelf to a high of more than 580 birds/day in June 2011 within the near-shore, coastal waters of the northern Gulf.

Persistence of avian carcasses on sandy beaches and marsh edges in the northern Gulf of Mexico.

As part of the natural resource damage assessment for the Deepwater Horizon oil spill, a mathematical model was used to estimate the total number of bird carcasses deposited on shorelines based on the number of carcasses collected and adjustment factors such as detection probability and carcass persistence. Studies of carcass persistence occurred along sandy beaches and marsh edges in the northern Gulf of Mexico to obtain site-specific inputs for the model. We estimated persistence rates for these habitat types and evaluated the influence on persistence of carcass size, location of the carcass on the beach, dominant vegetation type in the marsh, carcass distance into marsh vegetation, and length of time a carcass was stranded on a shoreline. The length of time stranded had the greatest influence on persistence in both habitat types, with persistence initially relatively low and increasing logarithmically. Carcass size and position were weakly influential on sandy beaches. Carcass size had stronger influences along marsh edges, and marsh habitat type also affected persistence. We found evidence of a positive relationship between distance into the marsh and persistence during the first 24 h after carcass deployment.

Chronic PAH exposures and associated declines in fish health indices observed for ten grouper species in the Gulf of Mexico.

Ten grouper species grouper (n = 584) were collected throughout the Gulf of Mexico (GoM) from 2011 through 2017 to provide information on hepatobiliary polycyclic aromatic hydrocarbon (PAH) concentrations in the aftermath of the Deepwater Horizon (DWH) oil spill. Liver and bile samples were analyzed for PAHs and their metabolites using triple quadrupole mass spectrometry (GC/MS/MS) and high-performance liquid chromatography with fluorescence detection (HPLC-F), respectively. Data were compared among species and sub-regions of the GoM to understand spatiotemporal exposure dynamics in these economically and ecologically important species. Significant differences in the composition and concentrations of PAHs were detected spatially, over time and by species. The West Florida Shelf, Cuba coast and the Yucatan Shelf had a greater proportion of the pyrogenic PAHs in their livers than the other regions likely due to non-oil industry related sources (e.g., marine vessel traffic) in the regional composition profiles. Mean liver PAH concentrations were highest in the north central region of the GoM where DWH occurred. Biliary PAH concentrations and health indicator biometrics initially decrease during the first three years following the DWH oil spill but significantly increased thereafter. Increased exposures are likely explained by the resuspension of residual DWH oil as well as continued inputs from natural (e.g., seeps) sources and other anthropogenically derived sources (e.g., riverine runoff, other oil spills, and leaking oil and gas infrastructure). The increasing trend in PAH concentrations in the bile and liver of grouper species in the north central region of the GoM post-DWH suggest continued chronic exposures, however the critical stage at which permanent, irreparable damage may occur is unknown. Long-term monitoring of PAH levels and associated fish health biomarkers is necessary to evaluate impacts of chronic exposures, particularly in regions subject to intensive oil extraction activities.

Petroleum hydrocarbons in semipermeable membrane devices deployed in the Northern Gulf of Mexico and Florida keys following the Deepwater Horizon incident.

The Deepwater Horizon (DWH) oil spill from April to July of 2010 contaminated Gulf of Mexico waters through release of an estimated 4.1 × 106 barrels of oil. Beginning in June of 2010, semipermeable membrane devices (SPMDs) were deployed near areas with sensitive marine habitats (Alabama Alps and Western Shelf) potentially exposed to that oil. Elevated TPAH50 concentrations, flux rates and similarity of histograms and diagnostic ratios for polycyclic aromatic hydrocarbons (PAH) from SPMDs to weathered floating oil collected during the DWH spill indicates the Alabama Alps habitats were affected. While not affected by oil from the DWH spill, the temporal pattern of PAH contamination of SPMDs deployed near the Western Shelf between July 2010 and March 2011 could indicate prevailing currents affected contaminant transport to the Western Shelf Area (East and West Flower Garden, Sonnier, and Stetson Banks) from non-DWH sources, including oil and gas exploration, shipping, and Mississippi River effluent.

Spatial contrasts in hepatic and biliary PAHs in Tilefish (Lopholatilus chamaeleonticeps) throughout the Gulf of Mexico, with comparison to the Northwest Atlantic.

A multinational demersal longline survey was conducted on the Gulf of Mexico continental shelf over the years 2015 and 2016 to generate a Gulf-wide baseline of polycyclic aromatic hydrocarbon (PAH) concentrations in demersal fishes. Tilefish (Lopholatilus chamaeleonticeps) were sampled in all regions of the Gulf of Mexico for biometrics, bile, and liver. Tilefish liver was also obtained from surveys in the northwest Atlantic Ocean for comparison. Liver tissues (n = 305) were analyzed for PAHs and select alkylated homologs using QuEChERS extractions and gas chromatography tandem mass spectrometry. Bile samples (n = 225) were analyzed for biliary PAH metabolites using high-performance liquid chromatography with fluorescence detection. Spatial comparisons indicate the highest levels of PAH exposure and hepatic accumulation in the north central Gulf of Mexico, with decreasing concentrations moving from the north central Gulf counterclockwise, and an increase on the Yucatán Shelf. Hepatic PAH concentrations were similar between the Gulf of Mexico and the northwest Atlantic, however, Tilefish from the northwest Atlantic had higher concentrations and more frequent detection of carcinogenic high molecular weight PAHs. Overall, results demonstrate that PAH pollution was ubiquitous within the study regions, with recent exposure and hepatic accumulation observed in Tilefish from both the Gulf of Mexico and northwest Atlantic.

Dispersant Enhances Hydrocarbon Degradation and Alters the Structure of Metabolically Active Microbial Communities in Shallow Seawater From the Northeastern Gulf of Mexico.

Dispersant application is a primary emergency oil spill response strategy and yet the efficacy and unintended consequences of this approach in marine ecosystems remain controversial. To address these uncertainties, ex situ incubations were conducted to quantify the impact of dispersant on petroleum hydrocarbon (PHC) biodegradation rates and microbial community structure at as close as realistically possible to approximated in situ conditions [2 ppm v/v oil with or without dispersant, at a dispersant to oil ratio (DOR) of 1:15] in surface seawater. Biodegradation rates were not substantially affected by dispersant application at low mixing conditions, while under completely dispersed conditions, biodegradation was substantially enhanced, decreasing the overall half-life of total PHC compounds from 15.4 to 8.8 days. While microbial respiration and growth were not substantially altered by dispersant treatment, RNA analysis revealed that dispersant application resulted in pronounced changes to the composition of metabolically active microbial communities, and the abundance of nitrogen-fixing prokaryotes, as determined by qPCR of nitrogenase (nifH) genes, showed a large increase. While the Gammaproteobacteria were enriched in all treatments, the Betaproteobacteria and different families of Alphaproteobacteria predominated in the oil and dispersant treatment, respectively. Results show that mixing conditions regulate the efficacy of dispersant application in an oil slick, and the quantitative increase in the nitrogen-fixing microbial community indicates a selection pressure for nitrogen fixation in response to a readily biodegradable, nitrogen-poor substrate.

Associations Between Chronic Exposure to Polycyclic Aromatic Hydrocarbons and Health Indices in Gulf of Mexico Tilefish (Lopholatilus chamaeleonticeps) Post Deepwater Horizon.

A time series of polycyclic aromatic hydrocarbon (PAH) data was collected for Gulf of Mexico demersal fishes in the years following the Deepwater Horizon oil spill (2012-2017). Tilefish (Lopholatilus chamaeleonticeps) were sampled via demersal longline at repeat stations in the northern Gulf of Mexico in 2012 to 2015 and 2017. Bile samples (n = 256) were analyzed via high-performance liquid chromatography with fluorescence detection for PAH metabolites as a biomarker of exposure to PAHs. Liver tissues (n = 230) were analyzed for accumulation of PAHs and alkylated homologs via quick, easy, cheap, effective, rugged, and safe (QuEChERS) extractions and gas chromatography-tandem mass spectrometry quantification. Over the 6-yr time series, exposure to petrogenic PAHs increased by an average of 178%, correlating with an average 22% decline in Fulton's condition factor. The decline in Fulton's condition factor was positively correlated with a 53% decline in percentage of liver lipid. There was no accumulation of PAHs in liver tissue over time. Together, these results suggest that increasing and chronic PAH exposure and metabolism may be taxing the energy budgets of tilefish, particularly adult females, with potentially negative impacts on fitness. Environ Toxicol Chem 2019;38:2659-2671. © 2019 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals, Inc. on behalf of SETAC.

Oil Spills and Human Health: Contributions of the Gulf of Mexico Research Initiative.

The Gulf of Mexico Research Initiative (GoMRI) was established in 2010 with $500 million in funding provided by British Petroleum over a 10-year period to support research on the impacts of the Deepwater Horizon oil spill and recovery. Contributions of the GoMRI program to date focused on human health are presented in more than 32 peer-reviewed papers published between 2011 and May 2019. Primary findings from review of these papers are (i) the large quantity of dispersants used in the oil cleanup have been associated with human health concerns, including through obesogenicity, toxicity, and illnesses from aerosolization of the agents; (ii) oil contamination has been associated with potential for increases in harmful algal blooms and numbers of pathogenic Vibrio bacteria in oil-impacted waters; and (iii) members of Gulf communities who are heavily reliant upon natural resources for their livelihoods were found to be vulnerable to high levels of life disruptions and institutional distrust. Positive correlations include a finding that a high level of community attachment was beneficial for recovery. Actions taken to improve disaster response and reduce stress-associated health effects could lessen negative impacts of similar disasters in the future. Furthermore, GoMRI has supported annual conferences beginning in 2013 at which informative human health-related presentations have been made. Based on this review, it is recommended that the Oil Pollution Act of 1990 be updated to include enhanced funding for oil spill impacts to human health.

Chemical characterization of natural and anthropogenic-derived oil residues on Gulf of Mexico beaches.

Oil residues originating from the Deepwater Horizon (DWH) incident persist on Gulf of Mexico beaches alongside oil from offshore industrial activity, natural seepage, and asphalt from parking lots and roads. To determine the primary differences in the chemical composition of these oil residues, a variety of samples were collected from beaches from Florida to Alabama over a two-year period from 2015 to 2017. Bulk chemical characteristics of the oil residues were examined via gas chromatography with flame ionization detection (GC-FID) and mass spectrometry (GC-MS), as well as thin layer chromatography with flame ionization detection (TLC-FID), and attenuated total reflectance Fourier transform infrared spectroscopy (ATR FT-IR). These bulk chemical analyses revealed features unique to the different sample types, expanding our understanding of the chemical composition and variability of persistent oil residues, and providing a means to detect and monitor their long-term fate in the coastal environment.

Applying spatiotemporal models to monitoring data to quantify fish population responses to the Deepwater Horizon oil spill in the Gulf of Mexico.

Quantifying the impacts of disturbances such as oil spills on marine species can be challenging. Natural environmental variability, human responses to the disturbance (e.g., fisheries closures), the complex life histories of the species being monitored, and limited pre-spill data can make detection of effects of oil spills difficult. Using long-term monitoring data from the state of Louisiana (USA), we applied novel spatiotemporal approaches to identify anomalies in species occurrence and catch rates. We included covariates (salinity, temperature, turbidity) to help isolate unusual events. While some species showed evidence of unlikely temporal anomalies in occurrence or catch rates, we found that the majority of the observed anomalies were also before the Deepwater Horizon event. Several species-gear combinations suggested upticks in the spatial variability immediately following the spill, but most species indicated no trend. Across species-gear combinations, there was no clear evidence for synchronous or asynchronous responses in occurrence or catch rates across sites following the spill. Our results are in general agreement to other analyses of monitoring data that detected small impacts, but in contrast to recent results from ecological modeling that showed much larger effects of the oil spill on fish and shellfish.

Corrigendum: The Effect of Hydrostatic Pressure on Enrichments of Hydrocarbon Degrading Microbes From the Gulf of Mexico Following the Deepwater Horizon Oil Spill.

[This corrects the article DOI: 10.3389/fmicb.2018.00808.].