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Cetaceans have a long history of commitment to a fully aquatic lifestyle that extends back to the Eocene. Extant species have evolved a spectacular array of adaptations in conjunction with their deployment into a diverse array of aquatic habitats. Sensory systems are among those that have experienced radical transformations in the evolutionary history of this clade. In the case of vision, previous studies have demonstrated important changes in the genes encoding rod opsin (RH1), short-wavelength sensitive opsin 1 (SWS1), and long-wavelength sensitive opsin (LWS) in selected cetaceans, but have not examined the full complement of opsin genes across the complete range of cetacean families. Here, we report protein-coding sequences for RH1 and both color opsin genes (SWS1, LWS) from representatives of all extant cetacean families. We examine competing hypotheses pertaining to the timing of blue shifts in RH1 relative to SWS1 inactivation in the early history of Cetacea, and we test the hypothesis that some cetaceans are rod monochomats. Molecular evolutionary analyses contradict the \"coastal\" hypothesis, wherein SWS1 was pseudogenized in the common ancestor of Cetacea, and instead suggest that RH1 was blue-shifted in the common ancestor of Cetacea before SWS1 was independently knocked out in baleen whales (Mysticeti) and in toothed whales (Odontoceti). Further, molecular evidence implies that LWS was inactivated convergently on at least five occasions in Cetacea: (1) Balaenidae (bowhead and right whales), (2) Balaenopteroidea (rorquals plus gray whale), (3) Mesoplodon bidens (Sowerby's beaked whale), (4) Physeter macrocephalus (giant sperm whale), and (5) Kogia breviceps (pygmy sperm whale). All of these cetaceans are known to dive to depths of at least 100 m where the underwater light field is dim and dominated by blue light. The knockout of both SWS1 and LWS in multiple cetacean lineages renders these taxa rod monochromats, a condition previously unknown among mammalian species.

The time and energetic costs of behavioral responses to incidental and experimental sonar exposures, as well as control stimuli, were quantified using hidden state analysis of time series of acoustic and movement data recorded by tags (DTAG) attached to 12 sperm whales (Physeter macrocephalus) using suction cups. Behavioral state transition modeling showed that tagged whales switched to a non-foraging, non-resting state during both experimental transmissions of low-frequency active sonar from an approaching vessel (LFAS; 1–2 kHz, source level 214 dB re 1 μPa m, four tag records) and playbacks of potential predator (killer whale, Orcinus orca) sounds broadcast at naturally occurring sound levels as a positive control from a drifting boat (five tag records). Time spent in foraging states and the probability of prey capture attempts were reduced during these two types of exposures with little change in overall locomotion activity, suggesting an effect on energy intake with no immediate compensation. Whales switched to the active non-foraging state over received sound pressure levels of 131–165 dB re 1 μPa during LFAS exposure. In contrast, no changes in foraging behavior were detected in response to experimental negative controls (no-sonar ship approach or noise control playback) or to experimental medium-frequency active sonar exposures (MFAS; 6–7 kHz, source level 199 re 1 μPa m, received sound pressure level [SPL] = 73–158 dB re 1 μPa). Similarly, there was no reduction in foraging effort for three whales exposed to incidental, unidentified 4.7–5.1 kHz sonar signals received at lower levels (SPL = 89–133 dB re 1 μPa). These results demonstrate that similar to predation risk, exposure to sonar can affect functional behaviors, and indicate that increased perception of risk with higher source level or lower frequency may modulate how sperm whales respond to anthropogenic sound.

The iron-limited Southern Ocean plays an important role in regulating atmospheric CO2 levels. Marine mammal respiration has been proposed to decrease the efficiency of the Southern Ocean biological pump by returning photosynthetically fixed carbon to the atmosphere. Here, we show that by consuming prey at depth and defecating iron-rich liquid faeces into the photic zone, sperm whales (Physeter macrocephalus) instead stimulate new primary production and carbon export to the deep ocean. We estimate that Southern Ocean sperm whales defecate 50 tonnes of iron into the photic zone each year. Molar ratios of Cexport ∶Feadded determined during natural ocean fertilization events are used to estimate the amount of carbon exported to the deep ocean in response to the iron defecated by sperm whales. We find that Southern Ocean sperm whales stimulate the export of 4 × 105 tonnes of carbon per year to the deep ocean and respire only 2 × 105 tonnes of carbon per year. By enhancing new primary production, the populations of 12 000 sperm whales in the Southern Ocean act as a carbon sink, removing 2 × 105 tonnes more carbon from the atmosphere than they add during respiration. The ability of the Southern Ocean to act as a carbon sink may have been diminished by large-scale removal of sperm whales during industrial whaling.

Sperm whales with the killer instinct Modern sperm whales have the biggest bite of any tetrapod, but their teeth are relatively small and restricted to the lower jaw, and they feed by suction. The discovery of large teeth in the fossil record had suggested, however, that predatory sperm whales once existed, similar in habit to today's killer whales, but much larger. Suggestion becomes reality with the discovery of the fossil teeth and jaws of a predatory sperm whale from the Middle Miocene of Peru, almost as large as a modern sperm whale but with a three-metre head and jaws full of teeth. This beast was probably able to prey upon large marine vertebrates — perhaps the medium-sized baleen whales abundant around 12 million years ago — in a way similar to the modern killer whale. Modern sperm whales have relatively small teeth and feed by suction, but the discovery of large teeth in the fossil record suggests that raptorial sperm whales once existed. Here the authors report the discovery of the teeth and jaws of a fossil raptorial sperm whale from the Middle Miocene of Peru, almost as large as a modern sperm whale but with a three-metre head and jaws full of teeth, some 36cm long. The modern giant sperm whale Physeter macrocephalus , one of the largest known predators, preys upon cephalopods at great depths 1 , 2 . Lacking a functional upper dentition, it relies on suction for catching its prey 3 ; in contrast, several smaller Miocene sperm whales (Physeteroidea) have been interpreted as raptorial (versus suction) feeders 4 , 5 , analogous to the modern killer whale Orcinus orca . Whereas very large physeteroid teeth have been discovered in various Miocene localities, associated diagnostic cranial remains have not been found so far 6 , 7 , 8 . Here we report the discovery of a new giant sperm whale from the Middle Miocene of Peru (approximately 12–13 million years ago), Leviathan melvillei , described on the basis of a skull with teeth and mandible. With a 3-m-long head, very large upper and lower teeth (maximum diameter and length of 12 cm and greater than 36 cm, respectively), robust jaws and a temporal fossa considerably larger than in Physeter , this stem physeteroid represents one of the largest raptorial predators and, to our knowledge, the biggest tetrapod bite ever found. The appearance of gigantic raptorial sperm whales in the fossil record coincides with a phase of diversification and size-range increase of the baleen-bearing mysticetes in the Miocene. We propose that Leviathan fed mostly on high-energy content medium-size baleen whales. As a top predator, together with the contemporaneous giant shark Carcharocles megalodon , it probably had a profound impact on the structuring of Miocene marine communities. The development of a vast supracranial basin in Leviathan , extending on the rostrum as in Physeter , might indicate the presence of an enlarged spermaceti organ in the former that is not associated with deep diving or obligatory suction feeding.

Sperm whalePhyseter macrocephalushabitat preferences are still poorly understood in the Mediterranean, despite the population being classified as ‘Endangered’ by the IUCN. Techniques to make the best use of multiple data sources are important in improving this situation. This work provides a detailed evaluation of sperm whale distribution and habitat use around the Balearic Islands using a novel analytical framework that combines transect and follow data while accounting for any autocorrelation present. During dedicated research cruises (2003 to 2008), sperm whales were located by listening at regular intervals along a search track and subsequently followed acoustically. Sperm whales were encountered 56 times and followed for periods ranging from a few hours to 3 d. Logistic Generalized Additive Models were used to model the probability of whale presence across the study area as a function of environmental variables, and Generalized Estimating Equations were used to account for autocorrelation. The results suggest that sperm whales do not use the region uniformly and that topography plays a key role in shaping their distribution. Moreover, solitary individuals were found to use the habitat differently from groups. This segregation appeared to be driven by water temperature and might reflect different needs or intraspecific competition. By shedding light on sperm whale habitat preference in such a critical area, this study represents an important step towards the implementation of conservation measures for this population.

We used generalized additive models (GAMs) as exploratory habitat models for describing the distribution of 2 deep-diving species, Cuvier's beaked whale Ziphius cavirostris Cuvier, 1823 and sperm whale Physeter catodon Linnaeus, 1758, in the Pelagos Sanctuary (northwestern Mediterranean). We analyzed data collected from research surveys and whalewatching activities during summer months from 2004 to 2007. The dataset encompassed 147 Cuvier's beaked whale sightings and 52 sperm whale sightings. We defined and applied a post hoc workflow to the data, to minimize false absence bias arising from the unique ecology of the species and the lack of a dedicated sampling design. We calculated a novel topographic predictor, distance from the canyon axis, as a covariate for use in the habitat model. Given the complex topography of the area, the analysis was performed on a highresolution spatial grid (1 km). Our methods allowed effective use of the nondedicated sampling dataset for building habitat models of elusive and cryptic species (Cuvier's beaked whale final model sensitivity = 0.88 and specificity = 0.84; sperm whale final model sensitivity = 0.65 and specificity = 0.77). The GAM results confirmed the preference for submarine canyons for both species and also highlighted the importance of the deeper portion of the Ligurian basin, especially for Cuvier's beaked whale. Habitat overlap nevertheless is resolved by a well-defined spatial partitioning of the area, with sperm whale occupying the western part and Cuvier's beaked whale the central and eastern parts.

Sperm whale (Physeter macrocephalus) populations were expected to rebuild following the end of commercial whaling. We document the decline of the population in the eastern Caribbean by tracing demographic changes of well-studied social units. We address hypotheses that, over a ten-year period of dedicated effort (2005-2015), unit size, numbers of calves and/or calving rates have each declined. Across 16 units, the number of adults decreased in 12 units, increased in two, and showed no change in two. The number of adults per unit decreased at -0.195 individuals/yr (95% CI: -0.080 to -0.310; P = 0.001). The number of calves also declined, but the decline was not significant. This negative trend of -4.5% per year in unit size started in about 2010, with numbers being fairly stable until then. There are several natural and anthropogenic threats, but no well-substantiated cause for the decline.

Predators make foraging decisions based upon sensory information about resource availability, but little is known about how large, air-breathing predators collect and use such information to maximize energy returns when foraging in the deep sea. Here, we used archival tags to study how echolocating sperm whales (Physeter macrocephalus) use their long-range sensory capabilities to guide foraging in a deep-water habitat consisting of multiple, depth-segregated prey layers. Sperm whales employ a directed search behaviour by modulating their overall sonar sampling with the intention to exploit a particular prey layer. They forage opportunistically during some descents while actively adjusting their acoustic gaze to sequentially track different prey layers. While foraging within patches, sperm whales adjust their clicking rate both to search new water volumes as they turn and to match the prey distribution. This strategy increases information flow and suggests that sperm whales can perform auditory stream segregation of multiple targets when echolocating. Such flexibility in sampling tactics in concert with long-range sensing capabilities apparently allow sperm whales to efficiently locate and access prey resources in vast, heterogeneous, deep water habitats.

The sperm whale (Physeter macrocephalus) emits a typical short acoustic signal, defined as a \"click\", almost continuously while diving. It is produced in different time patterns to acoustically explore the environment and communicate with conspecifics. Each emitted click has a multi-pulse structure, resulting from the production of the sound within the sperm whale's head. A Stable Inter Pulse Interval (Stable IPI) can be identified among the pulses that compose a single click. Applying specific algorithms, the measurement of this interval provides useful information to assess the total length of the animal recorded. In January 2005, a cabled hydrophone array was deployed at a depth of 2,100 m in the Central Mediterranean Sea, 25 km offshore Catania (Ionian Sea). The acoustic antenna, named ONDE (Ocean noise Detection Experiment), was in operation until November 2006. ONDE provided real time acoustic data used to perform Passive Acoustic Monitoring (PAM) of cetacean sound emissions. In this work, an innovative approach was applied to automatically measure the Stable IPI of the clicks, performing a cepstrum analysis to the energy (square amplitude) of the signals. About 2,100 five-minute recordings were processed to study the size distribution of the sperm whales detected during the ONDE long term deep-sea acoustic monitoring. Stable IPIs were measured in the range between 2.1 ms and 6.4 ms. The equations of Gordon (1991) and of Growcott (2011) were used to convert the IPIs into measures of size. The results revealed that the sperm whales recorded were distributed in length from about 7.5 m to 14 m. The size category most represented was from 9 m to 12 m (adult females or juvenile males) and specimens longer than 14 m (old males) seemed to be absent.

It is hypothesized that the primary function of permanent social relationships among female sperm whales (Physeter macrocephalus) is to provide allomothers for calves at the surface while mothers make foraging dives. In order to investigate how reciprocity of allocare within units of sperm whales facilitates group living, we constructed weighted social networks based on yearly matrices of associations (2005–2010) and correlated them across years, through changes in age and social role, to study changes in social relationships within seven sperm whale units. Pairs of association matrices from sequential years showed a greater positive correlation than expected by chance, but as the time lag increased, the correlation coefficients decreased. Over all units considered, calves had high values for all measured network statistics, while mothers had intermediate values for most of the measures, but high values for connectedness and affinity. Mothers showed sharp drops in strength and connectedness in the first year of their new calves' lives. These broad patterns appear to be consistent across units. Calves appeared to be significant nodes in the network of the social unit, and thus provide quantitative support for the theory in which communal care acts as the evolutionary force behind group formation in this species.