The influence of biopsy site and pregnancy on stable isotope ratios in humpback whale skin.

RATIONALE: Stable isotope analysis (SIA) of free-swimming mysticetes using biopsies is often limited in sample size and uses only one sample per individual, failing to capture both intra-individual variability and the influence of demographic and physiological factors on isotope ratios. METHODS: We applied SIA of δ13C and δ15N to humpback whale (Megaptera novaeangliae) biopsies taken during the foraging season along the western Antarctic Peninsula to quantify intra-individual variation from repeatedly sampled individuals, as well as to determine the effect of biopsy collection site, sex, and pregnancy on isotope ratios. RESULTS: There was substantial variability in δ13C from multiple biopsies taken from the same individuals, though δ15N was much more consistent. Side of the body (left versus right) and biopsy location (dorsal, anterior, ventral, and posterior) did marginally affect the isotopic composition of δ15N but not δ13C. Pregnancy had a significant effect on both δ13C and δ15N, where pregnant females were depleted in both when compared to non-pregnant females and males. CONCLUSIONS: These results indicate that isotopic signatures are influenced by multiple endogenous and exogenous factors and emphasize value in accounting for intra-individual variability and pregnancy status within a sampled population. Placed within an ecological context, the endogenous variability in δ13C observed here may be informative for future isotopic analyses.

Bellwethers of change: population modelling of North Pacific humpback whales from 2002 through 2021 reveals shift from recovery to climate response.

For the 40 years after the end of commercial whaling in 1976, humpback whale populations in the North Pacific Ocean exhibited a prolonged period of recovery. Using mark-recapture methods on the largest individual photo-identification dataset ever assembled for a cetacean, we estimated annual ocean-basin-wide abundance for the species from 2002 through 2021. Trends in annual estimates describe strong post-whaling era population recovery from 16 875 (± 5955) in 2002 to a peak abundance estimate of 33 488 (± 4455) in 2012. An apparent 20% decline from 2012 to 2021, 33 488 (± 4455) to 26 662 (± 4192), suggests the population abruptly reached carrying capacity due to loss of prey resources. This was particularly evident for humpback whales wintering in Hawai'i, where, by 2021, estimated abundance had declined by 34% from a peak in 2013, down to abundance levels previously seen in 2006, and contrasted to an absence of decline in Mainland Mexico breeding humpbacks. The strongest marine heatwave recorded globally to date during the 2014-2016 period appeared to have altered the course of species recovery, with enduring effects. Extending this time series will allow humpback whales to serve as an indicator species for the ecosystem in the face of a changing climate.

A collaborative and near-comprehensive North Pacific humpback whale photo-ID dataset.

We present an ocean-basin-scale dataset that includes tail fluke photographic identification (photo-ID) and encounter data for most living individual humpback whales (Megaptera novaeangliae) in the North Pacific Ocean. The dataset was built through a broad collaboration combining 39 separate curated photo-ID catalogs, supplemented with community science data. Data from throughout the North Pacific were aggregated into 13 regions, including six breeding regions, six feeding regions, and one migratory corridor. All images were compared with minimal pre-processing using a recently developed image recognition algorithm based on machine learning through artificial intelligence; this system is capable of rapidly detecting matches between individuals with an estimated 97-99% accuracy. For the 2001-2021 study period, a total of 27,956 unique individuals were documented in 157,350 encounters. Each individual was encountered, on average, in 5.6 sampling periods (i.e., breeding and feeding seasons), with an annual average of 87% of whales encountered in more than one season. The combined dataset and image recognition tool represents a living and accessible resource for collaborative, basin-wide studies of a keystone marine mammal in a time of rapid ecological change.

Detection probability and density estimation of fin whales by a Seaglider.

A single-hydrophone ocean glider was deployed within a cabled hydrophone array to demonstrate a framework for estimating population density of fin whales (Balaenoptera physalus) from a passive acoustic glider. The array was used to estimate tracks of acoustically active whales. These tracks became detection trials to model the detection function for glider-recorded 360-s windows containing fin whale 20-Hz pulses using a generalized additive model. Detection probability was dependent on both horizontal distance and low-frequency glider flow noise. At the median 40-Hz spectral level of 97 dB re 1 µPa2/Hz, detection probability was near one at horizontal distance zero with an effective detection radius of 17.1 km [coefficient of variation (CV) = 0.13]. Using estimates of acoustic availability and acoustically active group size from tagged and tracked fin whales, respectively, density of fin whales was estimated as 1.8 whales per 1000 km2 (CV = 0.55). A plot sampling density estimate for the same area and time, estimated from array data alone, was 1.3 whales per 1000 km2 (CV = 0.51). While the presented density estimates are from a small demonstration experiment and should be used with caution, the framework presented here advances our understanding of the potential use of gliders for cetacean density estimation.

Runs of homozygosity in killer whale genomes provide a global record of demographic histories.

Runs of homozygosity (ROH) occur when offspring inherit haplotypes that are identical by descent from each parent. Length distributions of ROH are informative about population history; specifically, the probability of inbreeding mediated by mating system and/or population demography. Here, we investigated whether variation in killer whale (Orcinus orca) demographic history is reflected in genome-wide heterozygosity and ROH length distributions, using a global data set of 26 genomes representative of geographic and ecotypic variation in this species, and two F1 admixed individuals with Pacific-Atlantic parentage. We first reconstructed demographic history for each population as changes in effective population size through time using the pairwise sequential Markovian coalescent (PSMC) method. We found a subset of populations declined in effective population size during the Late Pleistocene, while others had more stable demography. Genomes inferred to have undergone ancestral declines in effective population size, were autozygous at hundreds of short ROH (<1 Mb), reflecting high background relatedness due to coalescence of haplotypes deep within the pedigree. In contrast, longer and therefore younger ROH (>1.5 Mb) were found in low latitude populations, and populations of known conservation concern. These include a Scottish killer whale, for which 37.8% of the autosomes were comprised of ROH >1.5 Mb in length. The fate of this population, in which only two adult males have been sighted in the past five years, and zero fecundity over the last two decades, may be inextricably linked to its demographic history and consequential inbreeding depression.

Recommended snapshot length for acoustic point-transect surveys of intermittently available Cuvier's beaked whales.

Acoustic point-transect distance-sampling surveys have recently been used to estimate the density of beaked whales. Typically, the fraction of short time "snapshots" with detected beaked whales is used in this calculation. Beaked whale echolocation pulses are only intermittently available, which may affect the best choice of snapshot length. The effect of snapshot length on density estimation for Cuvier's beaked whale (Ziphius cavirostris) is investigated by sub-setting continuous recordings from drifting hydrophones deployed off southern and central California. Snapshot lengths from 20 s to 20 min are superimposed on the time series of detected beaked whale echolocation pulses, and the components of the density estimation equation are estimated for each snapshot length. The fraction of snapshots with detections, the effective area surveyed, and the snapshot detection probability all increase with snapshot length. Due to compensatory changes in these three components, density estimates show very little dependence on snapshot length. Within the range we examined, 1-2 min snapshots are recommended to avoid the potential bias caused by animal movement during the snapshot period and to maximize the sample size for estimating the effective area surveyed.

Acoustic detection range and population density of Cuvier's beaked whales estimated from near-surface hydrophones.

The population density of Cuvier's beaked whales is estimated acoustically with drifting near-surface hydrophone recorders in the Catalina Basin. Three empirical approaches (trial-based, distance-sampling, and spatially explicit capture-recapture) are used to estimate the probability of detecting the echolocation pulses as a function of range. These detection functions are used with two point-transect methods (snapshot and dive-cue) to estimate density. Measurement errors result in a small range of density estimates (3.9-5.4 whales per 1000 km2). Use of multiple approaches and methods allows comparison of the required information and assumptions of each. The distance-sampling approach with snapshot-based density estimates has the most stringent assumptions but would be the easiest to implement for large scale surveys of beaked whale density. Alternative approaches to estimating detection functions help validate this approach. The dive cue method of density estimation has promise, but additional work is needed to understand the potential bias caused by animal movement during a dive. Empirical methods are a viable alternative to the theoretical acoustic modeling approaches that have been used previously to estimate beaked whale density.

Time-Resolved Cost Analysis of Natural Gas Power Plant Conversion to Bioenergy with Carbon Capture and Storage to Support Net-Zero Emissions.

The ongoing construction of natural gas combined cycle (NGCC) power plants is incompatible with a transition to global net-zero greenhouse gas emissions. This work evaluates the emission pricing and technology costs required to convert an existing NGCC power plant to a biomethane-based bioenergy with carbon capture and storage (BECCS) system. The conversion was evaluated using techno-economic analysis and time-resolved life cycle assessment. At current technology costs, carbon dioxide equivalent emission prices of $142 and $215 per metric ton are required to allow a BECCS conversion to compete with normal operation or shutdown, respectively, of an existing NGCC power plant. These results show further technological development must occur in parallel with emission pricing to make BECCS viable. If mid-range emission pricing estimates are implemented ($25-$105 per metric ton), BECCS capital cost targets range from $1434 to $2098 per kW of capacity, while operational costs range from $32 to $51 per MWh of electricity produced to enable conversion. These findings indicate that operational costs associated with fuel consumption and production must be significantly reduced to make a BECCS conversion viable, even with emission pricing. All data and methods of this work have been made publicly available in an open-source model.

Performance evaluation of cetacean species distribution models developed using generalized additive models and boosted regression trees.

Species distribution models (SDMs) are important management tools for highly mobile marine species because they provide spatially and temporally explicit information on animal distribution. Two prevalent modeling frameworks used to develop SDMs for marine species are generalized additive models (GAMs) and boosted regression trees (BRTs), but comparative studies have rarely been conducted; most rely on presence-only data; and few have explored how features such as species distribution characteristics affect model performance. Since the majority of marine species BRTs have been used to predict habitat suitability, we first compared BRTs to GAMs that used presence/absence as the response variable. We then compared results from these habitat suitability models to GAMs that predict species density (animals per km2) because density models built with a subset of the data used here have previously received extensive validation. We compared both the explanatory power (i.e., model goodness of fit) and predictive power (i.e., performance on a novel dataset) of the GAMs and BRTs for a taxonomically diverse suite of cetacean species using a robust set of systematic survey data (1991-2014) within the California Current Ecosystem. Both BRTs and GAMs were successful at describing overall distribution patterns throughout the study area for the majority of species considered, but when predicting on novel data, the density GAMs exhibited substantially greater predictive power than both the presence/absence GAMs and BRTs, likely due to both the different response variables and fitting algorithms. Our results provide an improved understanding of some of the strengths and limitations of models developed using these two methods. These results can be used by modelers developing SDMs and resource managers tasked with the spatial management of marine species to determine the best modeling technique for their question of interest.

Detection and classification of narrow-band high frequency echolocation clicks from drifting recorders.

In the California Current off the United States West Coast, there are three offshore cetacean species that produce narrow-band high frequency (NBHF) echolocation pulses: Dall's porpoise (Phocoenoides dalli) and two species of Kogia. NBHF pulses exist in a highly specialized acoustic niche thought to be outside the hearing range of killer whales and other potential mammal-eating odontocetes. Very little is known about the dwarf and pygmy sperm whales (K. sima and K. breviceps), including their NBHF pulse characteristics. This paper presents a multivariate clustering method using data from unmanned drifting acoustic recorders and visually verified porpoise recordings to discriminate between probable porpoise and Kogia clicks. Using density clustering, this study finds three distinct clusters whose geographic distributions are consistent with the known habitat range for Kogia and Dall's porpoise. A Random Forest classification model correctly assigned 97% of the clicks to their cluster. Visually verified Dall's porpoise clicks from towed hydrophones were strongly associated with one of the clusters, while a second cluster tended to be outside the geographic range of Dall's porpoise and unlike the Dall's porpoise cluster. These clicks, presumed to be made by Kogia, exhibited greater spectral variance than previous Kogia echolocation studies. It is possible that the structure of Kogia NBHF pulses may not be as stereotypical as previously described.

Decline towards extinction of Mexico's vaquita porpoise (Phocoena sinus).

The vaquita (Phocoena sinus) is a small porpoise endemic to Mexico. It is listed by IUCN as Critically Endangered because of unsustainable levels of bycatch in gillnets. The population has been monitored with passive acoustic detectors every summer from 2011 to 2018; here we report results for 2017 and 2018. We combine the acoustic trends with an independent estimate of population size from 2015, and visual observations of at least seven animals in 2017 and six in 2018. Despite adoption of an emergency gillnet ban in May 2015, the estimated rate of decline remains extremely high: 48% decline in 2017 (95% Bayesian credible interval (CRI) 78% decline to 9% increase) and 47% in 2018 (95% CRI 80% decline to 13% increase). Estimated total population decline since 2011 is 98.6%, with greater than 99% probability the decline is greater than 33% yr-1. We estimate fewer than 19 vaquitas remained as of summer 2018 (posterior mean 9, median 8, 95% CRI 6-19). From March 2016 to March 2019, 10 dead vaquitas killed in gillnets were found. The ongoing presence of illegal gillnets despite the emergency ban continues to drive the vaquita towards extinction. Immediate management action is required if the species is to be saved.

Killer whale genomes reveal a complex history of recurrent admixture and vicariance.

Reconstruction of the demographic and evolutionary history of populations assuming a consensus tree-like relationship can mask more complex scenarios, which are prevalent in nature. An emerging genomic toolset, which has been most comprehensively harnessed in the reconstruction of human evolutionary history, enables molecular ecologists to elucidate complex population histories. Killer whales have limited extrinsic barriers to dispersal and have radiated globally, and are therefore a good candidate model for the application of such tools. Here, we analyse a global data set of killer whale genomes in a rare attempt to elucidate global population structure in a nonhuman species. We identify a pattern of genetic homogenisation at lower latitudes and the greatest differentiation at high latitudes, even between currently sympatric lineages. The processes underlying the major axis of structure include high drift at the edge of species' range, likely associated with founder effects and allelic surfing during postglacial range expansion. Divergence between Antarctic and non-Antarctic lineages is further driven by ancestry segments with up to four-fold older coalescence time than the genome-wide average; relicts of a previous vicariance during an earlier glacial cycle. Our study further underpins that episodic gene flow is ubiquitous in natural populations, and can occur across great distances and after substantial periods of isolation between populations. Thus, understanding the evolutionary history of a species requires comprehensive geographic sampling and genome-wide data to sample the variation in ancestry within individuals.

Time Value of Greenhouse Gas Emissions in Life Cycle Assessment and Techno-Economic Analysis.

Life cycle assessment is a fundamental tool used to evaluate the environmental impact of products. Standard life cycle assessment methodology ignores the impact of greenhouse gases relative to when they are emitted. In this paper, we present a method for leveraging the social cost of greenhouse gases to account for the temporal impacts of emissions in life cycle assessment and techno-economics. To demonstrate, we use this method to analyze the present value of the monetized impacts of emissions across multiple electricity generation technologies. Results show that accounting for time increases the present value across all but one of the technologies considered. Carbon intensive technologies show the highest increase, with coal rising between 26% and 62% depending on social cost scenario. Additionally, we demonstrate a second method that combines temporally resolved greenhouse gas emissions with techno-economic analysis. Considering temporal impacts of emissions within techno-economic analysis increases the levelized cost of electricity (LCOE) across all technologies considered. Carbon intensive technologies increase significantly, with the LCOE from coal rising between 37% and 263% depending on the social cost scenario. The proposed methods show that temporal resolution in life cycle assessment is critical for comparing the monetized impacts of greenhouse gas emissions across technologies.

Diving behavior of Cuvier's beaked whales inferred from three-dimensional acoustic localization and tracking using a nested array of drifting hydrophone recorders.

Echolocation pulses from Cuvier's beaked whales are used to track the whales' three-dimensional diving behavior in the Catalina Basin, California. In 2016, five 2-element vertical hydrophone arrays were suspended from the surface and drifted at ∼100-m depth. Cuvier's beaked whale pulses were identified, and vertical detection angles were estimated from time-differences-of-arrival of either direct-path signals received on two hydrophones or direct-path and surface-reflected signals received on the same hydrophone. A Bayesian state-space model is developed to track the diving behavior. The model is fit to these detection angle estimates from at least four of the drifting vertical arrays. Results show that the beaked whales were producing echolocation pulses and are presumed to be foraging at a mean depth of 967 m (standard deviation = 112 m), approximately 300 m above the bottom in this basin. Some whales spent at least some time at or near the bottom. Average swim speed was 1.2 m s-1, but swim direction varied during a dive. The average net horizontal speed was 0.6 m s-1. Results are similar to those obtained from previous tagging studies of this species. These methods may allow expansion of dive studies to other whale species that are difficult to tag.

Last call: Passive acoustic monitoring shows continued rapid decline of critically endangered vaquita.

The vaquita is a critically endangered species of porpoise. It produces echolocation clicks, making it a good candidate for passive acoustic monitoring. A systematic grid of sensors has been deployed for 3 months annually since 2011; results from 2016 are reported here. Statistical models (to compensate for non-uniform data loss) show an overall decline in the acoustic detection rate between 2015 and 2016 of 49% (95% credible interval 82% decline to 8% increase), and total decline between 2011 and 2016 of over 90%. Assuming the acoustic detection rate is proportional to population size, approximately 30 vaquita (95% credible interval 8-96) remained in November 2016.

Precision and bias in estimating detection distances for beaked whale echolocation clicks using a two-element vertical hydrophone array.

Detection distances are critical for cetacean density and abundance estimation using distance sampling methods. Data from a drifting buoy system consisting of an autonomous recorder and a two-element vertical hydrophone array at ∼100-m depth are used to evaluate three methods for estimating the horizontal distance (range) to beaked whales making echolocation clicks. The precision in estimating time-differences-of-arrival (TDOA) for direct- and surface-reflected-path clicks is estimated empirically using repeated measures over short time periods. A Teager-Kaiser energy detector is used to improve estimates of TDOA for surface-reflected signals. Simulations show that array tilt in the direction of the source cannot be reliably estimated given this array geometry and these measurements of TDOA error, which means that range cannot be reliably estimated. If array tilt can be reduced to less than 0.5°, range can be reliably estimated up to ∼3000 m. If array depth is increased to 200 m and array tilt is less than 1°, range can be reliably estimated up to ∼5000 m. Prior information on the depth of vocalizing beaked whales and estimates of declination angle can be used to precisely estimate range, but different analytical methods are required to avoid bias and to treat distributions of depth probabilistically.

Acoustic differentiation of Shiho- and Naisa-type short-finned pilot whales in the Pacific Ocean.

Divergence in acoustic signals used by different populations of marine mammals can be caused by a variety of environmental, hereditary, or social factors, and can indicate isolation between those populations. Two types of genetically and morphologically distinct short-finned pilot whales, called the Naisa- and Shiho-types when first described off Japan, have been identified in the Pacific Ocean. Acoustic differentiation between these types would support their designation as sub-species or species, and improve the understanding of their distribution in areas where genetic samples are difficult to obtain. Calls from two regions representing the two types were analyzed using 24 recordings from Hawai'i (Naisa-type) and 12 recordings from the eastern Pacific Ocean (Shiho-type). Calls from the two types were significantly differentiated in median start frequency, frequency range, and duration, and were significantly differentiated in the cumulative distribution of start frequency, frequency range, and duration. Gaussian mixture models were used to classify calls from the two different regions with 74% accuracy, which was significantly greater than chance. The results of these analyses indicate that the two types are acoustically distinct, which supports the hypothesis that the two types may be separate sub-species.

Using paired visual and passive acoustic surveys to estimate passive acoustic detection parameters for harbor porpoise abundance estimates.

Passive acoustic monitoring is a promising approach for monitoring long-term trends in harbor porpoise (Phocoena phocoena) abundance. Before passive acoustic monitoring can be implemented to estimate harbor porpoise abundance, information about the detectability of harbor porpoise is needed to convert recorded numbers of echolocation clicks to harbor porpoise densities. In the present study, paired data from a grid of nine passive acoustic click detectors (C-PODs, Chelonia Ltd., United Kingdom) and three days of simultaneous aerial line-transect visual surveys were collected over a 370 km2 study area. The focus of the study was estimating the effective detection area of the passive acoustic sensors, which was defined as the product of the sound production rate of individual animals and the area within which those sounds are detected by the passive acoustic sensors. Visually estimated porpoise densities were used as informative priors in a Bayesian model to solve for the effective detection area for individual harbor porpoises. This model-based approach resulted in a posterior distribution of the effective detection area of individual harbor porpoises consistent with previously published values. This technique is a viable alternative for estimating the effective detection area of passive acoustic sensors when other experimental approaches are not feasible.

Passive acoustic monitoring of the decline of Mexico's critically endangered vaquita.

The vaquita (Phocoena sinus) is the world's most endangered marine mammal with approximately 245 individuals remaining in 2008. This species of porpoise is endemic to the northern Gulf of California, Mexico, and historically the population has declined because of unsustainable bycatch in gillnets. An illegal gillnet fishery for an endangered fish, the totoaba (Totoaba macdonaldi), has recently resurged throughout the vaquita's range. The secretive but lucrative wildlife trade with China for totoaba swim bladders has probably increased vaquita bycatch mortality by an unknown amount. Precise population monitoring by visual surveys is difficult because vaquitas are inherently hard to see and have now become so rare that sighting rates are very low. However, their echolocation clicks can be identified readily on specialized acoustic detectors. Acoustic detections on an array of 46 moored detectors indicated vaquita acoustic activity declined by 80% between 2011 and 2015 in the central part of the species' range. Statistical models estimated an annual rate of decline of 34% (95% Bayesian credible interval -48% to -21%). Based on results from 2011 to 2014, the government of Mexico enacted and is enforcing an emergency 2-year ban on gillnets throughout the species' range to prevent extinction, at a cost of US$74 million to compensate fishers. Developing precise acoustic monitoring methods proved critical to exposing the severity of vaquitas' decline and emphasizes the need for continual monitoring to effectively manage critically endangered species.