Morphology of nares associated with stereo-olfaction in baleen whales.

The sensory mechanisms used by baleen whales (Mysticeti) for locating ephemeral, dense prey patches in vast marine habitats are poorly understood. Baleen whales have a functional olfactory system with paired rather than single blowholes (nares), potentially enabling stereo-olfaction. Dimethyl sulfide (DMS) is an odorous gas emitted by phytoplankton in response to grazing by zooplankton. Some seabirds use DMS to locate prey, but this ability has not been demonstrated in whales. For 14 extant species of baleen whale, nares morphometrics (imagery from unoccupied aerial systems, UAS) was related to published trophic level indices using Bayesian phylogenetic mixed modelling. A significant negative relationship was found between nares width and whale trophic level (ß = -0.08, lower 95% CI = -0.13, upper 95% CI = -0.03), corresponding with a 39% increase in nares width from highest to lowest trophic level. Thus, species with nasal morphology best suited to stereo-olfaction are more zooplanktivorous. These findings provide evidence that some baleen whale species may be able to localize odorants e.g. DMS. Our results help direct future behavioural trials of olfaction in baleen whales, by highlighting the most appropriate species to study. This is a research priority, given the potential for DMS-mediated plastic ingestion by whales.

Energy expenditure of southern right whales varies with body size, reproductive state and activity level.

Quantifying the energy expenditure of animals is critical to understanding the cost of anthropogenic disturbance relative to their overall energy requirements. We used novel drone focal follows (776 follows, 185 individuals) and aerial photogrammetry (5372 measurements, 791 individuals) to measure the respiration rate and body condition loss of southern right whales (Eubalaena australis) on a breeding ground in Australia. Respiration rates were converted to oxygen consumption rate and field metabolic rate (FMR) using published bioenergetic models. The intra-seasonal loss in body condition of different reproductive classes (calves, juveniles, adults, pregnant and lactating females) was converted to blubber energy loss and total energy expenditure (TEE). Using these two metrics, we tested the effects of body size, reproductive state and activity level on right whale energy expenditure. Respiration rates and mass-specific FMR decreased exponentially with an increase in body size, as expected based on allometric scaling. FMR increased curvilinearly with an increase in swim speed, probably as a result of increased drag and increased locomotion costs. Respiration rates and FMR were 44% higher for pregnant and lactating females compared with those of adults, suggesting significant costs of fetal maintenance and milk production, respectively. The estimated FMR of adults based on their respiration rates corresponded well with the estimated TEE based on body condition loss. The rate of decline in body condition of pregnant and lactating females was considerably higher than expected based on respiration rates, which probably reflects the milk energy transfer from mothers to calves, which is not reflected in their FMR.

Fetal growth, birth size and energetic cost of gestation in southern right whales.

The cost of reproduction greatly affects a species' life history strategy. Baleen whales exhibit some of the fastest offspring growth rates in the animal kingdom. We quantified the energetic cost of gestation for southern right whales (Eubalaena australis) by combining whaling catch records of pregnant females with photogrammetry data on southern right whale mothers and calves from two breeding grounds in Argentina and Australia. The relationship between calf birth size and maternal length was determined from repeated measurements of individual females before and after giving birth. Fetal growth was determined from generalized linear models fitted to fetal length data from whaling operations between 1961 and 1967. Fetal length was converted to volume and mass, using the volume-to-length relationship of newborn southern right whales calves, and published tissue composition and energy content estimates. Fetal maintenance costs (heat of gestation) and the energy content of the placenta were predicted from published relationships and added to the fetal growth cost to calculate the total cost of gestation. Our findings showed that fetal growth rates and birth size increased linearly with maternal length, with calves being born at ∼35% maternal length. Fetal length increased curvilinearly through gestation, which resulted in an exponential increase in fetal volume and mass. Consequently, the cost of gestation was very low during the first (0.1% of total cost) and second trimester (4.9%), but increased rapidly during the last trimester (95.0%). The heat of gestation incurred the highest cost for pregnant females (73.8%), followed by fetal growth (21.2%) and the placental energy content (5.0%). KEY POINTS: Baleen whales exhibit some of the fastest fetal growth rates in the animal kingdom. Despite this, the energetic cost of gestation is largely unknown, as well as the influence of maternal body size on fetal growth rates and calf birth sizes. We combined historical whaling records and drone photogrammetry data to determine fetal growth rates and birth sizes in southern right whales (Eubalaena australis), from which we estimated the cost of gestation. Calf birth size, and consequent fetal growth rates, increased positively with maternal body size. The cost of gestation was negligible for southern right whale females during the first two trimesters, but increased rapidly during the last trimester. These results show that late gestation incurs a significant cost for baleen whale females, and needs to be accounted for in bioenergetic models.

Scaling of maneuvering performance in baleen whales: larger whales outperform expectations.

Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.

Acoustic crypsis in southern right whale mother-calf pairs: infrequent, low-output calls to avoid predation?

Southern right whales (Eubalaena australis) invest substantial amounts of energy in their calves, while facing the risk of having them predated upon by eavesdropping killer whales (Orcinus orca). We tested the hypothesis that southern right whale mother-calf pairs employ acoustic crypsis to reduce acoustic detectability by such predators. Specifically, we deployed multi-sensor DTAGs on nine lactating whales for a total of 62.9 h in a Western Australian breeding ground, and used a SoundTrap to estimate the concomitant acoustic background noise. Vocalisations were recorded at low rates of <10 calls h-1 (1 call per dive) and at low received levels between 123±8 and 134±10 dB re. 1 µPa RMS depending on call type. We conclude that such acoustic crypsis in southern right whales and other baleen whales decreases the risk of alerting potential predators and hence jeopardizing a substantial energetic investment by the mother.

El Niño Southern Oscillation influences the abundance and movements of a marine top predator in coastal waters.

Large-scale climate modes such as El Niño Southern Oscillation (ENSO) influence population dynamics in many species, including marine top predators. However, few quantitative studies have investigated the influence of large-scale variability on resident marine top predator populations. We examined the effect of climate variability on the abundance and temporary emigration of a resident bottlenose dolphin (Tursiops aduncus) population off Bunbury, Western Australia (WA). This population has been studied intensively over six consecutive years (2007-2013), yielding a robust dataset that captures seasonal variations in both abundance and movement patterns. In WA, ENSO affects the strength of the Leeuwin Current (LC), the dominant oceanographic feature in the region. The strength and variability of the LC affects marine ecosystems and distribution of top predator prey. We investigated the relationship between dolphin abundance and ENSO, Southern Annular Mode, austral season, rainfall, sea surface salinity and sea surface temperature (SST). Linear models indicated that dolphin abundance was significantly affected by ENSO, and that the magnitude of the effect was dependent upon season. Dolphin abundance was lowest during winter 2009, when dolphins had high temporary emigration rates out of the study area. This coincided with the single El Niño event that occurred throughout the study period. Coupled with this event, there was a negative anomaly in SST and an above average rainfall. These conditions may have affected the distribution of dolphin prey, resulting in the temporary emigration of dolphins out of the study area in search of adequate prey. This study demonstrated the local effects of large-scale climatic variations on the short-term response of a resident, coastal delphinid species. With a projected global increase in frequency and intensity of extreme climatic events, resident marine top predators may not only have to contend with increasing coastal anthropogenic activities, but also have to adapt to large-scale climatic changes.

Sexual Dimorphism and Geographic Variation in Dorsal Fin Features of Australian Humpback Dolphins, Sousa sahulensis.

Determining the sex of free-ranging cetaceans can be challenging. Sexual dimorphism among external features may allow inferences on sex, but such patterns may be difficult to detect and are often confounded by age and geographic variation. Dorsal fin images of 107 female and 54 male Australian humpback dolphins, Sousa sahulensis, from Western Australia (WA) and Queensland (QLD) were used to investigate sex, age and geographic differences in colouration, height/length quotient and number of notches. Adult males exhibited more dorsal fin notches (p<0.001) and a significantly greater loss of pigmentation on the upper half of their dorsal fins (p<0.001) than did adult females. These differences likely reflect that males experience a higher frequency and/or intensity of intraspecific aggression than females. In QLD, heavily spotted dorsal fins were more frequent among females than males (p<0.001). Logistic regression analyses revealed that dorsal fin spotting and loss of pigmentation on the upper half of the dorsal fin provided the best model parameters for predicting the sex of sampled adults, with 97% accuracy. This technique offers a rapid, non-invasive method for predicting sex in Australian humpback dolphins, which could potentially be applied to populations throughout their range. In contrast to adults, presumed immature animals showed little or no loss of pigmentation or spotting; however, the rate of development of these features remains unknown. There were pronounced differences between QLD and WA in the intensity of spotting on dorsal fins and the extent of pigmentation loss around the posterior insertion and trailing edge of the dorsal fin. While based on a limited sample size, these geographic differences may have conservation implications in terms of population subdivision and should be investigated further.

Passive acoustic monitoring of coastally associated Hawaiian spinner dolphins, Stenella longirostris, ground-truthed through visual surveys.

Effective decision making to protect coastally associated dolphins relies on monitoring the presence of animals in areas that are critical to their survival. Hawaiian spinner dolphins forage at night and rest during the day in shallow bays. Due to their predictable presence, they are targeted by dolphin-tourism. In this study, comparisons of presence were made between passive acoustic monitoring (PAM) and vessel-based visual surveys in Hawaiian spinner dolphin resting bays. DSG-Ocean passive acoustic recording devices were deployed in four bays along the Kona Coast of Hawai'i Island between January 8, 2011 and August 30, 2012. The devices sampled at 80 kHz, making 30-s recordings every four minutes. Overall, dolphins were acoustically detected on 37.1% to 89.6% of recording days depending on the bay. Vessel-based visual surveys overlapped with the PAM surveys on 202 days across the four bays. No significant differences were found between visual and acoustic detections suggesting acoustic surveys can be used as a proxy for visual surveys. Given the need to monitor dolphin presence across sites, PAM is the most suitable and efficient tool for monitoring long-term presence/absence. Concomitant photo-identification surveys are necessary to address changes in abundance over time.

Echolocation parameters of Australian humpback dolphins (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus) in the wild.

Echolocation is a key sensory modality for toothed whale orientation, navigation, and foraging. However, a more comparative understanding of the biosonar properties of toothed whales is necessary to understand behavioral and evolutionary adaptions. To address this, two free-ranging sympatric delphinid species, Australian humpback dolphins (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus), were studied. Biosonar clicks from both species were recorded within the same stretch of coastal habitat in Exmouth Gulf, Western Australia, using a vertical seven element hydrophone array. S. sahulensis used biosonar clicks with a mean source level of 199 ± 3 dB re 1 µPa peak-peak (pp), mean centroid frequency of 106 ± 11 kHz, and emitted at interclick intervals (ICIs) of 79 ± 33 ms. These parameters were similar to click parameters of sympatric T. aduncus, characterized by mean source levels of 204 ± 4 dB re 1 µPa pp, centroid frequency of 112 ± 9 kHz, and ICIs of 73 ± 29 ms. These properties are comparable to those of other similar sized delphinids and suggest that biosonar parameters are independent of sympatric delphinids and possibly driven by body size. The dynamic biosonar behavior of these delphinids may have, consequently, allowed for adaptations to local environments through high levels of control over sonar beam properties.

Cetacean morbillivirus in coastal Indo-Pacific bottlenose dolphins, Western Australia.

Cetacean morbillivirus (CeMV) has caused several epizootics in multiple species of cetaceans globally and is an emerging disease among cetaceans in Australia. We detected CeMV in 2 stranded coastal Indo-Pacific bottlenose dolphins (Tursiops aduncus) in Western Australia. Preliminary phylogenetic data suggest that this virus variant is divergent from known strains.

Cultural transmission of tool use combined with habitat specializations leads to fine-scale genetic structure in bottlenose dolphins.

Socially learned behaviours leading to genetic population structure have rarely been described outside humans. Here, we provide evidence of fine-scale genetic structure that has probably arisen based on socially transmitted behaviours in bottlenose dolphins (Tursiops sp.) in western Shark Bay, Western Australia. We argue that vertical social transmission in different habitats has led to significant geographical genetic structure of mitochondrial DNA (mtDNA) haplotypes. Dolphins with mtDNA haplotypes E or F are found predominantly in deep (more than 10 m) channel habitat, while dolphins with a third haplotype (H) are found predominantly in shallow habitat (less than 10 m), indicating a strong haplotype-habitat correlation. Some dolphins in the deep habitat engage in a foraging strategy using tools. These 'sponging' dolphins are members of one matriline, carrying haplotype E. This pattern is consistent with what had been demonstrated previously at another research site in Shark Bay, where vertical social transmission of sponging had been shown using multiple lines of evidence. Using an individual-based model, we found support that in western Shark Bay, socially transmitted specializations may have led to the observed genetic structure. The reported genetic structure appears to present an example of cultural hitchhiking of mtDNA haplotypes on socially transmitted foraging strategies, suggesting that, as in humans, genetic structure can be shaped through cultural transmission.

Quantifying the abundance and survival rates of island-associated spinner dolphins using a multi-state open robust design model.

Spinner dolphins (Stenella longirostris subsp.) occupy the nearshore waters of several Hawaiian Islands. Due to their constrained behavioral pattern and genetic isolation, they are vulnerable to anthropogenic threats. Their occurrence and behavior are well-described, yet a lack of data on their abundance and survival rates hinders optimal conservation action. Using design-based photo-identification surveys, this study estimated the abundance, apparent survival, and emigration of spinner dolphins off the Wai'anae Coast of O'ahu through multi-state open robust design (MSORD) and POPAN modelling. Eight seasonal field seasons, (two winter, spring, summer, and autumn) each comprised of six surveys of the study area, were completed during two consecutive years. Seasonal abundance estimates derived from the best fitting model ranged from 140 (± 36.8 SE, 95% CI 84-232) to 373 (± 60.0, 95% CI 273-509) individuals and were lowest during winter seasons. The MSORD estimated a survival rate of 0.95 (± 0.02 SE) and a Markovian pattern of temporary emigration. POPAN modelling estimated a super-population size of 633 (± 78 SE, 95% CI 492-798), reflecting the total number of individual dolphins that used the study area during the entire study period. Additional research on circum- and inter-island dolphin movements around and between O'ahu and the Maui Nui region may shed light on both seasonal movement patterns and overall abundance for the O'ahu/4-Islands stock. This work represents the first systematic mark-recapture effort to assess the abundance and survival rates of these highly exposed dolphins, providing valuable insights for conservation and management.

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.

Social and genetic interactions drive fitness variation in a free-living dolphin population.

The evolutionary forces that drive fitness variation in species are of considerable interest. Despite this, the relative importance and interactions of genetic and social factors involved in the evolution of fitness traits in wild mammalian populations are largely unknown. To date, a few studies have demonstrated that fitness might be influenced by either social factors or genes in natural populations, but none have explored how the combined effect of social and genetic parameters might interact to influence fitness. Drawing from a long-term study of wild bottlenose dolphins in the eastern gulf of Shark Bay, Western Australia, we present a unique approach to understanding these interactions. Our study shows that female calving success depends on both genetic inheritance and social bonds. Moreover, we demonstrate that interactions between social and genetic factors also influence female fitness. Therefore, our study represents a major methodological advance, and provides critical insights into the interplay of genetic and social parameters of fitness.

Quantifying the age structure of free-ranging delphinid populations: Testing the accuracy of Unoccupied Aerial System photogrammetry.

Understanding the population health status of long-lived and slow-reproducing species is critical for their management. However, it can take decades with traditional monitoring techniques to detect population-level changes in demographic parameters. Early detection of the effects of environmental and anthropogenic stressors on vital rates would aid in forecasting changes in population dynamics and therefore inform management efforts. Changes in vital rates strongly correlate with deviations in population growth, highlighting the need for novel approaches that can provide early warning signs of population decline (e.g., changes in age structure). We tested a novel and frequentist approach, using Unoccupied Aerial System (UAS) photogrammetry, to assess the population age structure of small delphinids. First, we measured the precision and accuracy of UAS photogrammetry in estimating total body length (TL) of trained bottlenose dolphins (Tursiops truncatus). Using a log-transformed linear model, we estimated TL using the blowhole to dorsal fin distance (BHDF) for surfacing animals. To test the performance of UAS photogrammetry to age-classify individuals, we then used length measurements from a 35-year dataset from a free-ranging bottlenose dolphin community to simulate UAS estimates of BHDF and TL. We tested five age classifiers and determined where young individuals (<10 years) were assigned when misclassified. Finally, we tested whether UAS-simulated BHDF only or the associated TL estimates provided better classifications. TL of surfacing dolphins was overestimated by 3.3% ±3.1% based on UAS-estimated BHDF. Our age classifiers performed best in predicting age-class when using broader and fewer (two and three) age-class bins with ~80% and ~72% assignment performance, respectively. Overall, 72.5%-93% of the individuals were correctly classified within 2 years of their actual age-class bin. Similar classification performances were obtained using both proxies. UAS photogrammetry is a non-invasive, inexpensive, and effective method to estimate TL and age-class of free-swimming dolphins. UAS photogrammetry can facilitate the detection of early signs of population changes, which can provide important insights for timely management decisions.

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.

Estimated communication range and energetic cost of bottlenose dolphin whistles in a tropical habitat.

Bottlenose dolphins (Tursiops sp.) depend on frequency-modulated whistles for many aspects of their social behavior, including group cohesion and recognition of familiar individuals. Vocalization amplitude and frequency influences communication range and may be shaped by many ecological and physiological factors including energetic costs. Here, a calibrated GPS-synchronized hydrophone array was used to record the whistles of bottlenose dolphins in a tropical shallow-water environment with high ambient noise levels. Acoustic localization techniques were used to estimate the source levels and energy content of individual whistles. Bottlenose dolphins produced whistles with mean source levels of 146.7 ± 6.2 dB re. 1 µPa(RMS). These were lower than source levels estimated for a population inhabiting the quieter Moray Firth, indicating that dolphins do not necessarily compensate for the high noise levels found in noisy tropical habitats by increasing their source level. Combined with measured transmission loss and noise levels, these source levels provided estimated median communication ranges of 750 m and maximum communication ranges up to 5740 m. Whistles contained less than 17 mJ of acoustic energy, showing that the energetic cost of whistling is small compared to the high metabolic rate of these aquatic mammals, and unlikely to limit the vocal activity of toothed whales.

Residency, movement patterns, behavior and demographics of reef manta rays in Komodo National Park.

Background: The reef manta ray (Mobula alfredi) is a globally threatened species and an iconic tourist attraction for visitors to Indonesia's Komodo National Park (NP). In 2013, manta ray fishing was banned in Komodo NP and its surroundings, preceding the nationwide manta ray protection in 2014. Over a decade ago, a previous acoustic telemetry study demonstrated that reef manta rays had high fidelity to sites within the park, while more recent photo-identification data indicated that some individuals move up to 450 km elsewhere. Characterization of manta ray demographics, behavior, and a focused assessment on site use of popular tourism locations within the park is vital to assist the Komodo NP Management Authority formulate appropriate manta ray conservation and management policies. Methods: This study uses a long-term library (MantaMatcher.org) of photo-identification data collected by researchers and citizen scientists to investigate manta ray demographics and habitat use within the park at four sites frequented by tour operators: Cauldron, Karang Makassar, Mawan, and Manta Alley. Residency and movements of manta rays were investigated with maximum likelihood analyses and Markov movement models. Results: A total of 1,085 individual manta rays were identified from photographs dating from 2013 to 2018. In general, individual manta rays displayed a higher affinity to specific sites than others. The highest re-sighting probabilities came from the remote southern site, Manta Alley. Karang Makassar and Mawan are only ~5 km apart; however, manta rays displayed distinct site affinities. Exchange of individuals between Manta Alley and the two central sites (~35.5 km apart) occurred, particularly seasonally. More manta rays were recorded traveling from the south to the central area than vice versa. Female manta rays were more mobile than males. Similar demographic groups used Karang Makassar, Mawan, and Manta Alley for foraging, cleaning, cruising, or courtship activities. Conversely, a higher proportion of immature manta rays used the northern site, Cauldron, where foraging was commonly observed. Fishing gear-related injuries were noted on 56 individuals (~5%), and predatory injuries were present on 32 individuals (~3%). Tourism within the park increased from 2014 to 2017, with 34% more dive boats per survey at Karang Makassar and Mawan. Discussion: The Komodo NP contains several distinct critical habitats for manta rays that encompass all demographics and accommodate seasonal manta ray movements. While the present study has not examined population trends, it does provide foundational data for such work. Continued research into manta ray abundance, long-range movements, and identifying and protecting other critical aggregation areas within the region is integral to securing the species' recovery. We provide management recommendations to limit undue pressure on manta rays and their critical habitats from tourism.

Fast and Furious: Energetic Tradeoffs and Scaling of High-Speed Foraging in Rorqual Whales.

Although gigantic body size and obligate filter feeding mechanisms have evolved in multiple vertebrate lineages (mammals and fishes), intermittent ram (lunge) filter feeding is unique to a specific family of baleen whales: rorquals. Lunge feeding is a high cost, high benefit feeding mechanism that requires the integration of unsteady locomotion (i.e., accelerations and maneuvers); the impact of scale on the biomechanics and energetics of this foraging mode continues to be the subject of intense study. The goal of our investigation was to use a combination of multi-sensor tags paired with UAS footage to determine the impact of morphometrics such as body size on kinematic lunging parameters such as fluking timing, maximum lunging speed, and deceleration during the engulfment period for a range of species from minke to blue whales. Our results show that, in the case of krill-feeding lunges and regardless of size, animals exhibit a skewed gradient between powered and fully unpowered engulfment, with fluking generally ending at the point of both the maximum lunging speed and mouth opening. In all cases, the small amounts of propulsive thrust generated by the tail were unable to overcome the high drag forces experienced during engulfment. Assuming this thrust to be minimal, we predicted the minimum speed of lunging across scale. To minimize the energetic cost of lunge feeding, hydrodynamic theory predicts slower lunge feeding speeds regardless of body size, with a lower boundary set by the ability of the prey to avoid capture. We used empirical data to test this theory and instead found that maximum foraging speeds remain constant and high (∼4 m s-1) across body size, even as higher speeds result in lower foraging efficiency. Regardless, we found an increasing relationship between body size and this foraging efficiency, estimated as the ratio of energetic gain from prey to energetic cost. This trend held across timescales ranging from a single lunge to a single day and suggests that larger whales are capturing more prey-and more energy-at a lower cost.

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus).

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 µPa peak to peak, click durations of 8-72 µs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.