Comparative genomics provides insights into the aquatic adaptations of mammals.

The ancestors of marine mammals once roamed the land and independently committed to an aquatic lifestyle. These macroevolutionary transitions have intrigued scientists for centuries. Here, we generated high-quality genome assemblies of 17 marine mammals (11 cetaceans and six pinnipeds), including eight assemblies at the chromosome level. Incorporating previously published data, we reconstructed the marine mammal phylogeny and population histories and identified numerous idiosyncratic and convergent genomic variations that possibly contributed to the transition from land to water in marine mammal lineages. Genes associated with the formation of blubber (NFIA), vascular development (SEMA3E), and heat production by brown adipose tissue (UCP1) had unique changes that may contribute to marine mammal thermoregulation. We also observed many lineage-specific changes in the marine mammals, including genes associated with deep diving and navigation. Our study advances understanding of the timing, pattern, and molecular changes associated with the evolution of mammalian lineages adapting to aquatic life.

Echolocation clicks of free-ranging Indo-Pacific finless porpoises (Neophocaena phocaenoides) in Hainan waters.

The echolocation clicks of free-ranging Indo-Pacific finless porpoises (IPFPs, Neophocaena phocaenoides) have been rarely studied in the wild. This paper aims at describing the echolocation-click characteristics of IPFPs and examining whether IPFPs adapt their sonar system to the habitats in Hainan waters, China. The echolocation clicks were recorded using a 13 elements star-shaped array of hydrophones. A total of 65 on-axis clicks were identified and analyzed. IPFPs use echolocation clicks with a source level (SL) of 158 ± 9 dB re: 1 µPa peak-peak, mean peak, and centroid frequency of 134 ± 3 kHz, -3 dB bandwidth of 14 ± 2 kHz and produce at inter-click intervals of 104 ± 51 ms. The results relative to other porpoises show that finless porpoises in Hainan waters produce clicks with moderate SLs and high peak frequency. These results could be useful in detecting the presence and estimating the density of IPFPs during passive acoustic monitoring in the study area and serve to shed light on the interpopulation variation of click characteristics of finless porpoises as well.

Ranging pattern development of a declining delphinid population: A potential cascade effect of vessel activities.

It is generally accepted that vessel activity causes various behavioral responses of cetaceans and undermines individual fitness. Whether or how it can lead to a demographic response of populations remains rarely examined. In the northern Beibu Gulf, China, vessel activities have sharply increased in the past two decades, while abnormal demographic dynamics was recently noted for the resident Indo-Pacific humpback dolphins. The present study first examined the humpback dolphins' utilization distribution (UD) from 2003 to 2019. Habitat suitability was then modeled with the sighting data collected before the most recent population reduction. Finally, we tried to disentangle the anthropogenic driver of dolphin demography by cross-referring the spatiotemporal development of dolphins' UD, vessel activities, and habitat suitability. Our results showed that the dolphins' UD shrank substantially during the port expansion in the early 2010s, and we suggest that the consequential increase in vessel activities might impose extra marine stressors on the resident humpback dolphins. To reduce the boat interaction, the dolphins steadily shifted their core area to a less suitable area in the east during 2015-2017, when unnaturally low survivals were recorded. Afterward, the dolphin core area partially shifted back to the more suitable area in the west, which corresponded to the improving dolphin survival in 2018. Our finding suggested that the vessel activity may be responsible for the dolphin displacement, while staying in the less suitable area may further lead to a more severe and acute demographic consequence on the population. The underlying and indirect impact of vessel activities as disclosed by the present study is particularly important for port management, marine planning, and conservation practice regarding coastal cetaceans, especially for those resident and endangered populations inhabiting the urbanized coastal areas.

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.

Numerical-modeling-based investigation of sound transmission and reception in the short-finned pilot whale (Globicephala macrorhynchus).

The sound-transmission, beam-formation, and sound-reception processes of a short-finned pilot whale (Globicephala macrorhynchus) were investigated using computed tomography (CT) scanning and numerical simulation. The results showed that sound propagations in the forehead were modulated by the upper jaw, air components, and soft tissues, which attributed to the beam formation in the external acoustic field. These structures owned different acoustic impedance and formed a multiphasic sound transmission system that can modulate sounds into a beam. The reception pathways composed of the solid mandible and acoustic fats in the lower head conducted sounds into the tympano-periotic complex. In the simulations, sounds were emitted in the forehead transmission system and propagated into water to interrogate a steel cylinder. The resulting echoes can be interpreted from multiple perspectives, including amplitude, waveform, and spectrum, to obtain the acoustic cues of the steel cylinder. By taking the short-finned pilot whale as an example, this study provides meaningful information to further deepen our understanding of biosonar system operations, and may expand sound-reception theory in odontocetes.

Influence of acoustic habitat variation on Indo-Pacific humpback dolphin (Sousa chinensis) in shallow waters of Hainan Island, China.

The Indo-Pacific humpback dolphin (IPHD, Sousa chinensis) is a coastal species inhabiting tropical and warm-temperate waters. The presence of this vulnerable dolphin was recently discovered in shallow waters southwest of Hainan Island, China. The influence of the acoustic habitat on the distribution and behavior of IPHD was investigated using an array of passive acoustic platforms (n = 6) that spanned more than 100 km of coastline during a 75-day monitoring period. Its presence was assessed within 19 215 five-min recordings by classifying echolocation clicks using machine learning techniques. Spectrogram analysis was applied to further investigate the acoustic behavior of IPHD and to identify other prominent sound sources. The variation in the ambient noise levels was also measured to describe the spatiotemporal patterns of the acoustic habitat among the different sampling sites. Social and feeding sounds of IPHD (whistles and click-series of pulsed sounds) were identified together with other biological sources (finless porpoise, soniferous fishes, and snapping shrimps) and anthropogenic activities (ship noise, explosions, and sonars). Distribution, acoustic behavior, and habitat use of this nearshore dolphin species were strongly influenced by the abundance of soniferous fishes, and under similar conditions, the species was more acoustically active in locations with lower noise levels.

Echolocation signals of free-ranging pantropical spotted dolphins (Stenella attenuata) in the South China Sea.

Echolocation signals of free-ranging pantropical spotted dolphins (Stenella attenuata) in the western Pacific Ocean have not been studied much. This paper aims to describe the characteristics of echolocation signals of S. attenuata in the northern South China Sea. A six-arm star array with 13 hydrophones was used and a total of 131 on-axis clicks were identified to analyze the acoustic features of the echolocation signals of dolphins. The mean center frequency was 89 ± 13 kHz, with mean peak-to-peak sound source levels of 190 ± 6 dB re: 1 µPa @ 1 m. The mean -3 dB bandwidth and root-mean-square bandwidth were 62 ± 15 kHz and 26 ± 3 kHz, respectively, with mean -10 dB duration of 18 ± 4 µs and root-mean-square duration of 6 ± 2 µs. The results showed that click parameters of S. attenuata in the northern South China Sea are different from those of clicks of the species in Hawaii waters. The differences in click parameters may be due to both behavioral context and/or environmental adaptation of S. attenuata in different habitats.

Whistles emitted by Indo-Pacific humpback dolphins (Sousa chinensis) in Zhanjiang waters, China.

Whistles emitted by Indo-Pacific humpback dolphins in Zhanjiang waters, China, were collected by using autonomous acoustic recorders. A total of 529 whistles with clear contours and signal-to-noise ratio higher than 10 dB were extracted for analysis. The fundamental frequencies and durations of analyzed whistles were in ranges of 1785-21 675 Hz and 30-1973 ms, respectively. Six tonal types were identified: constant, downsweep, upsweep, concave, convex, and sine whistles. Constant type was the most dominant tonal type, accounting for 32.51% of all whistles, followed by sine type, accounting for 19.66% of all whistles. This paper examined 17 whistle parameters, which showed significant differences among the six tonal types. Whistles without inflections, gaps, and stairs accounted for 62.6%, 80.6%, and 68.6% of all whistles, respectively. Significant intraspecific differences in all duration and frequency parameters of dolphin whistles were found between this study and the study in Malaysia. Except for start frequency, maximum frequency and the number of harmonics, all whistle parameters showed significant differences between this study and the study conducted in Sanniang Bay, China. The intraspecific differences in vocalizations for this species may be related to macro-geographic and/or environmental variations among waters, suggesting a potential geographic isolation among populations of Indo-Pacific humpback dolphins.

Acoustic recordings of rough-toothed dolphin (Steno bredanensis) offshore Eastern Sicily (Mediterranean Sea).

Rough-toothed dolphin's abundance and distribution is largely unknown worldwide and evaluation of its conservation status in the Mediterranean Sea is necessary. A rough-toothed dolphin was sighted offshore Eastern Sicily (Mediterranean Sea) in July 2017 and acoustic data were acquired in the same area of Watkins, Tyack, Moore, and Notarbartolo di Sciara [(1987). Mar. Mamm. Sci. 3, 78-82]. An automatic detection algorithm was developed to identify the echolocation clicks recorded within both datasets and a recurrent inter-click interval value was identified during the new encounter. Distinctive whistle classes were also identified with similar contour shapes within both datasets.

Broadband ship noise and its potential impacts on Indo-Pacific humpback dolphins: Implications for conservation and management.

Ship noise pollution has raised considerable concerns among regulatory agencies and cetacean researchers worldwide. There is an urgent need to quantify ship noise in coastal areas and assess its potential biological impacts. In this study, underwater broadband noise from commercial ships in a critical habitat of Indo-Pacific humpback dolphins was recorded and analyzed. Data analysis indicated that the ship noise caused by the investigated commercial ships with an average length of 134 ± 81 m, traveling at 18.8 ± 2.5 km/h [mean ± standard deviation (SD), n = 21] comprises mid-to-high components with frequencies approaching and exceeding 100 kHz, and the ship noise could be sensed auditorily by Indo-Pacific humpback dolphins within most of their sensitive frequency range. The contributions of ship noise to ambient noise were highest in two third-octave bands with center frequencies of 8 and 50 kHz, which are within the sensitive hearing range of Indo-Pacific humpback dolphins and overlap the frequency of sounds that are biologically significant to the dolphins. It is estimated that ship noise in these third-octave bands can be auditorily sensed by and potentially affect the dolphins within 2290 ± 1172 m and 848 ± 358 m (mean ± SD, n = 21), respectively.

Acoustic occurrence detection of a newly recorded Indo-Pacific humpback dolphin population in waters southwest of Hainan Island, China.

In 2014, Indo-Pacific humpback dolphins were recorded for the first time in waters southwest of Hainan Island, China. In this paper, the temporal occurrence of Indo-Pacific humpback dolphins in this region was detected by stationary passive acoustic monitoring. During the 130-day observation period (from January to July 2016), 1969 click trains produced by Indo-Pacific humpback dolphins were identified, and 262 ten-minute recording bins contained echolocation click trains of dolphins, of which 70.9% were at night and 29.1% were during the day. A diurnal rhythm with a nighttime peak in acoustic detections was found. Passive acoustic detections indicated that the Indo-Pacific humpback dolphins frequently occurred in this area and were detected mainly at night. This information may be relevant to conservation efforts for these dolphins in the near future.

The influence of air-filled structures on wave propagation and beam formation of a pygmy sperm whale (Kogia breviceps) in horizontal and vertical planes.

The wave propagation, sound field, and transmission beam pattern of a pygmy sperm whale (Kogia breviceps) were investigated in both the horizontal and vertical planes. Results suggested that the signals obtained at both planes were similarly characterized with a high peak frequency and a relatively narrow bandwidth, close to the ones recorded from live animals. The sound beam measured outside the head in the vertical plane was narrower than that of the horizontal one. Cases with different combinations of air-filled structures in both planes were used to study the respective roles in controlling wave propagation and beam formation. The wave propagations and beam patterns in the horizontal and vertical planes elucidated the important reflection effect of the spermaceti and vocal chambers on sound waves, which was highly significant in forming intensive forward sound beams. The air-filled structures, the forehead soft tissues and skull structures formed wave guides in these two planes for emitted sounds to propagate forward.

The echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis).

While the transmission beam of odontocetes has been described in a number of studies, the majority of them that have measured the transmission beam in two dimensions were focused on captive animals. Within the current study, a dedicated cross hydrophone array with nine elements was used to investigate the echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins. A total of 265 on-axis clicks were analyzed, from which the apparent peak to peak source levels ranged between 168 to 207 dB (mean 184.5 dB ± 6.6 dB). The 3-dB beam width along the horizontal and vertical plane was 9.6° and 7.4°, respectively. Measured separately, the directivity index of the horizontal and vertical plane was 12.6 and 13.5 dB, respectively, and the overall directivity index (both planes combined) was 29.5 dB. The beam shape was slightly asymmetrical along the horizontal and vertical axis. Compared to other species, the characteristics of the transmitting beam of Indo-Pacific humpback dolphins were relatively close to the bottlenose dolphin (Tursiops truncatus), likely due to the similarity in the peak frequency and waveform of echolocation clicks and comparable body sizes of the two species.

Acoustic properties of a short-finned pilot whale head with insight into temperature influence on tissues' sound velocity.

Acoustic properties of odontocete head tissues, including sound velocity, density, and acoustic impedance, are important parameters to understand dynamics of its echolocation. In this paper, acoustic properties of head tissues from a freshly dead short-finned pilot whale (Globicephala macrorhynchus) were reconstructed using computed tomography (CT) and ultrasound. The animal's forehead soft tissues were cut into 188 ordered samples. Sound velocity, density, and acoustic impedance of each sample were either directly measured or calculated by formula, and Hounsfield Unit values (HUs) were obtained from CT scanning. According to relationships between HUs and sound velocity, HUs and density, as well as HUs and acoustic impedance, distributions of acoustic properties in the head were reconstructed. The inner core in the melon with low-sound velocity and low-density is an evidence for its potential function of sound focusing. The increase in acoustic impedance of forehead tissues from inner core to outer layer may be important for the acoustic impedance matching between the outer layer tissue and seawater. In addition, temperature dependence of sound velocity in soft tissues was also examined. The results provide a guide to the simulation of the sound emission of the short-finned pilot whale.

Likely Age-Related Hearing Loss (Presbycusis) in a Stranded Indo-Pacific Humpback Dolphin (Sousa chinensis).

The hearing of a stranded Indo-Pacific humpback dolphin (Sousa chinensis) in Zhuhai, China, was measured. The age of this animal was estimated to be ~40 years. The animal's hearing was measured using a noninvasive auditory evoked potential (AEP) method. The results showed that the high-frequency hearing cutoff frequency of the studied dolphin was ~30-40 kHz lower than that of a conspecific younger individual ~13 year old. The lower high-frequency hearing range in the older dolphin was explained as a likely result of age-related hearing loss (presbycusis).

In-air vocal repertoires of spotted seals, Phoca largha.

Spotted seals (Phoca largha) are thought to be less vocal than other phocids. However, acoustic communication behaviors of spotted seals have been reported several times. In this study, the vocal repertoires of spotted seals housed in Dalian Sun Aquarium, China were recorded and analyzed. The frequencies of the sounds made by the seals ranged from 139.3 to 2323.1 Hz, and the time durations lasted from 92.8 to 1208 ms, depending on age and gender (P < 0.01). The peak-to-peak sound source levels were 109-124 dB re 20µPa. In total, seven vocal types were identified: pup call, yearling call, bark, growl, grunt, moo, and throat guttural. The pups emitted sounds with high frequencies (F1: 972.4 ± 374.4 Hz, mean ± standard deviation) and medial time durations (564 ± 178 ms); when the pups grew older, the sounds became yearling calls, which had high frequencies with median (interquartile range) of 1198.0 (821.7-1385.5) Hz; and long time durations [902 (745-1080) ms]. The male adults emitted sounds with low frequencies [430.2 (388.2-486.7) Hz] and short time durations [334 (233-599) ms], while the female adults emitted sounds with medial frequencies [814.5 (592.6-1024.3) Hz] and medial time durations [531 (336-688) ms].

Echolocation signals of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis) in Sanniang Bay, China.

While the low-frequency communication sounds of Indo-Pacific humpback dolphins (Sousa chinensis) have been reported in a number of papers, the high-frequency echolocation signals of Sousa chinensis, especially those living in the wild, have been less studied. In the current study, echolocation signals of humpback dolphins were recorded in Sanniang Bay, Guangxi Province, China, using a cross-type hydrophone array with five elements. In total, 77 candidate on-axis clicks from 77 scans were selected for analysis. The results showed that the varied peak-to-peak source levels ranged from 177.1 to 207.3 dB, with an average of 187.7 dB re: 1 µPa. The mean peak frequency was 109.0 kHz with a -3-dB bandwidth of 50.3 kHz and 95% energy duration of 22 µs. The -3-dB bandwidth was much broader than the root mean square bandwidth and exhibited a bimodal distribution. The center frequency exhibited a positive relationship with the peak-to-peak source level. The clicks of the wild Indo-Pacific humpback dolphins were short-duration, broadband, ultrasonic pulses, similar to those produced by other whistling dolphins of similar body size. However, the click source levels of the Indo-Pacific humpback dolphin appear to be lower than those of other whistling dolphins.

Mid- to high-frequency noise from high-speed boats and its potential impacts on humpback dolphins.

The impact of noise made by vessels on marine animals has come under increased concern. However, most measurements on noise from vessels have only taken into account the low-frequency components. For cetaceans operating in the mid- and high-frequencies, such as the Indo-Pacific humpback dolphin (Sousa chinensis), mid- to high-frequency noise components may be of more concern, in terms of their potential impacts. In this study, noise made by a small high-speed boat was recorded using a broadband recording system in a dolphin watching area focusing on the effects on humpback dolphins in Sanniang Bay, China. The high-speed boat produced substantial mid- to high-frequency noise components with frequencies to >100 kHz, measured at three speeds: ∼40, 30, and 15 km/h. The noise from the boat raised the ambient noise levels from ∼5 to 47 decibels (dB) root-mean-square (rms) across frequency bands ranging from 1 to 125 kHz at a distance of 20 to 85 m, with louder levels recorded at higher speeds and at closer distances. To conclude, the noise produced by the small high-speed boat could be heard by Sousa chinensis and therefore potentially had adverse effects on the dolphins.

Acoustic property reconstruction of a pygmy sperm whale (Kogia breviceps) forehead based on computed tomography imaging.

Computed tomography (CT) imaging and sound experimental measurements were used to reconstruct the acoustic properties (density, velocity, and impedance) of the forehead tissues of a deceased pygmy sperm whale (Kogia breviceps). The forehead was segmented along the body axis and sectioned into cross section slices, which were further cut into sample pieces for measurements. Hounsfield units (HUs) of the corresponding measured pieces were obtained from CT scans, and regression analyses were conducted to investigate the linear relationships between the tissues' HUs and velocity, and HUs and density. The distributions of the acoustic properties of the head at axial, coronal, and sagittal cross sections were reconstructed, revealing that the nasal passage system was asymmetric and the cornucopia-shaped spermaceti organ was in the right nasal passage, surrounded by tissues and airsacs. A distinct dense theca was discovered in the posterior-dorsal area of the melon, which was characterized by low velocity in the inner core and high velocity in the outer region. Statistical analyses revealed significant differences in density, velocity, and acoustic impedance between all four structures, melon, spermaceti organ, muscle, and connective tissue (p < 0.001). The obtained acoustic properties of the forehead tissues provide important information for understanding the species' bioacoustic characteristics.