Niche overlap and diet composition of three sympatric coastal dolphin species in the southwest Atlantic Ocean

Sympatric species are expected to differ in ecological requirements to minimize niche overlap and avoid competition. Here we assess the trophic interactions among three coexisting dolphin species from southern Brazil: the franciscana dolphin (Pontoporia blainvillei), the Guiana dolphin (Sotalia guianensis), and the Lahille's bottlenose dolphin (Tursiops truncatus gephyreus). We evaluated temporal variation in carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen to examine potential dietary shifts resulting from increased fishing activity over the past three decades. We estimated the degree of niche overlap among these species and the contribution of potential prey sources to their diet. δ¹⁵N values were consistent among species and across years, while δ¹³C values increased for Guiana dolphins and decreased for bottlenose dolphins, suggesting changes in diet and/or foraging habitats through time. The similar δ¹³C and δ¹⁵N values and the high niche overlap between Guiana and bottlenose dolphins indicate that these species are primarily feeding on demersal prey. The franciscana diet is primarily composed of pelagic prey, resulting in a lower niche overlap in comparison with the other dolphin species. Our study provides further information about the foraging ecology of this unique dolphin community in southern Brazil with implications for its management and conservation.     

CETACEAN ABUNDANCE IN HAWAIIAN WATERS ESTIMATED FROM A SUMMER/FALL SURVEY IN 2002

Cetacean abundance is estimated for the U.S. Exclusive Economic Zone (EEZ) around the Hawaiian Islands based on a ship line‐transect survey from August to November, 2002. Sighting detection functions are estimated from this and other NOAA research surveys from 1986 to 2002 using a new, multiple‐covariate approach. Twenty‐four species were seen on this survey, including two species (Fraser's dolphin [Lagenodelphis hosei] and sei whale [Balaenoptera borealis]) that had not been previously documented to occur in Hawaiian waters. The most abundant large whales are sperm whales (Physeter macrocephalus) and Bryde's whales (Balaenoptera edeni). The most abundant delphinids are pilot whales (Globicephala macrorhynchus), rough‐toothed dolphins (Steno bredanensis), Fraser's dolphins, spotted dolphins (Stenella attenuata), and striped dolphins (Stenella coeruleoalba). Dwarf and pygmy sperm whales (Kogia sima and Kogia breviceps) and Cuvier's beaked whales (Ziphius cavirostris) are also estimated to be quite abundant. Some of the migratory baleen whales (fin whales [Balaenoptera physalus], sei whales, minke whales [B. acutorostrata], and humpback whales [Megaptera novaeangliae]) were seen only late in the survey. Abundance is estimated for 19 cetacean species. The overall density of cetaceans is low in the study area, especially for delphinids. The precision of density and abundance estimates is generally low for all species because of the low number of sightings.     

Differences in acoustic signals from Delphinids in the western North Atlantic and northern Gulf of Mexico

Whistle characteristics were quantitatively compared between both geographically separated and neighboring populations of Atlantic spotted dolphins (Stenella frontalis), bottlenose dolphins (Tursiops truncatus), and pilot whales (Globicephala spp.) in U.S. waters to evaluate if intraspecific acoustic differences exist between groups. We compared nine whistle characteristics between continental shelf and offshore Atlantic spotted dolphins in the western North Atlantic and between northern Gulf of Mexico and western North Atlantic bottlenose dolphins and pilot whales using discriminant analysis. Offshore Atlantic spotted dolphin whistles were significantly different (Hotelling's T², P= 0.0003) from continental shelf whistles in high frequency, bandwidth, duration, number of steps, and number of inflection points. Atlantic bottlenose dolphin whistles were significantly different (Hotelling's T², P < 0.0001) from those in the Gulf of Mexico in duration, number of steps, and number of inflection points. There was no significant difference between pilot whale whistles in the two basins. The whistle differences indicate acoustic divergence between groups in different areas that may arise from geographic isolation or habitat separation between neighboring but genetically distinct populations of dolphins. This study supports the premise that acoustic differences can be a tool to evaluate the ecological separation between marine mammal groups in field studies.     

Detection of specific antibodies to morbilliviruses, Brucella and Toxoplasma in the Black Sea dolphin Tursiops truncatus ponticus and the beluga whale Delphinapterus leucas from the Sea of Okhotsk in 2002–2007

The prevalence of antibodies to morbilliviruses, Brucella and Toxoplasma was studied in the Black Sea bottlenose dolphin Tursiops truncatus ponticus and the beluga whale Delphinapterus leucas from the Sea of Okhotsk. The blood serum of 74 dolphins and 147 beluga whales was tested in 2002–2007. Antibodies to morbilliviruses were detected in 15 (20.3%) bottlenose dolphins and 20 (13.6%) beluga whales. Antibodies to Brucella were detected in 17 (23.0%) bottlenose dolphins and 10 (6.8%) beluga whales. Toxoplasma-specific antibodies were detected in 39 (52.7%) bottlenose dolphins and 7 (4.8%) beluga whales. Some animals had antibodies to two, or even three, of the pathogens. A high level of incidence of the pathogens in the sea animals was found in the densely populated coastal areas with high economic development.     

Evidence for infanticide in bottlenose dolphins: an explanation for violent interactions with harbour porpoises?

Most harbour porpoises found dead on the north-east coast of Scotland show signs of attack by sympatric bottlenose dolphins, but the reason(s) for these violent interactions remain(s) unclear. Post-mortem examinations of stranded bottlenose dolphins indicate that five out of eight young calves from this same area were also killed by bottlenose dolphins. These data, together with direct observations of an aggressive interaction between an adult bottlenose dolphin and a dead bottlenose dolphin calf, provide strong evidence for infanticide in this population. The similarity in the size range of harbour porpoises and dolphin calves that showed signs of attack by bottlenose dolphins suggests that previously reported interspecific interactions could be related to this infanticidal behaviour. These findings appear to provide the first evidence of infanticide in cetaceans (whales, dolphins and porpoises). We suggest that infanticide must be considered as a factor shaping sociality in this and other species of cetaceans, and may have serious consequences for the viability of small populations.     

Food partitioning among three sympatric odontocetes (Grampus griseus,Lagenodelphis hosei,and Stenella attenuata)

Food is one of the most important dimensions of resource partitioning for species coexistence. In this study, we investigated the dietary composition and foraging habits of three sympatric odontocetes in order to identify their levels of food niche overlap and ecological separation. Stomach content analysis was performed on samples collected from carcasses confiscated by police or entangled in gill nets from 1994 to 2001, including 27 Risso's dolphins (GG) (Grampus griseus), 27 Fraser's dolphins (LH) (Lagenodelphis hosei), and 45 pantropical spotted dolphins (SA) (Stenella attenuata). GG consumed only cephalopods, with Enoploteuthis chunii accounting for 90.5% of total prey consumed, LH fed on mesopelagic fishes and cephalopods, dominated by hatchetfish, Polyipnus stereope (50.2%), and SA ate both mesopelagic and epipelagic preys, primarily fishes of Myctophum asperum (20.3%) and squids of E. chunii (25.8%). Among the three odontocetes, GG had the narrowest dietary niche width, while SA had the widest width. Both the niche overlap index and the analysis of similarities (ANOSIM) showed significant diet differentiation among these three dolphin species. The depth distribution of their principal prey items further suggests that LH feeds in the deepest waters while SA utilizes prey resources near surface.     

Incomplete reporting of whale, dolphin and porpoise 'bycatch' revealed by molecular monitoring of Korean markets

We report the results of molecular monitoring of 'whalemeat' markets in the Republic of (South) Korea based on nine systematic surveys from February 2003 to February 2005. As Korea has no programme of commercial or scientific whaling and there is a closure on the hunting of dolphins and porpoises, the only legal source of these products was assumed to be incidental fisheries mortalities ('bycatch') as reported by the government to the International Whaling Commission. Species identification of 357 products using mitochondrial DNA control region or cytochrome b sequences and the web-based programme DNA-surveillance revealed three species of baleen whales (North Pacific minke, common form Bryde's and humpback), three species of beaked whales (Cuvier's, Stejneger's and Blainville's), seven species of dolphins (short-finned pilot, false killer and killer whales; Risso's, bottlenose, common and Pacific white-sided dolphins) and two species of porpoises (harbour and finless). Comparison of market products with official records revealed a number of discrepancies. Of the eight species identified on the markets in 2003, three were not reported in official records for that year. Of the 11 species identified in 2004, five were not reported as bycatch, although one species, a humpback whale, was reported as 'stranded'. We also found significant inconsistencies in the expected frequencies of products from most species, including a large over-representation of finless porpoises and false killer whales. We suggest ways in which market surveys could be improved to provide better information on the magnitude of fisheries bycatch and other illegal, unregulated and unreported (IUU) exploitation of wildlife.     

Vocal activity of tropical dolphins is inhibited by the presence of killer whales,Orcinus orca

Research has suggested killer whale (Orcinus orca) predation may affect cetacean vocal behavior; however, few data exist to test this hypothesis. Data collected during 40,976 km of visual and acoustic shipboard surveys in the tropical Pacific Ocean, including 1,232 detections of 13 species, were examined to determine if changes in dolphin vocal activity could be attributed to the presence of killer whales. Generalized linear models and Random Forest analyses were used to test the hypothesis that dolphin vocal activity was related to the distance and time to the nearest killer whale sighting. Both results show that dolphin vocalizations were inversely correlated with the temporal proximity of killer whales (P < 0.05). Despite the relative rarity of killer whales in the tropics, they appear to influence vocal behavior of nearby dolphin schools. This disruption in communication may not significantly impact interactions necessary for survival in tropical waters where killer whale density is low. However, in temperate climates, where increased productivity supports a greater abundance of killer whales, this interruption in communication may have a greater impact. The lower incidence of whistling dolphins in temperate waters may be related to the greater abundance of killer whales in these areas.     

Hearing loss in stranded odontocete dolphins and whales

The causes of dolphin and whale stranding can often be difficult to determine. Because toothed whales rely on echolocation for orientation and feeding, hearing deficits could lead to stranding. We report on the results of auditory evoked potential measurements from eight species of odontocete cetaceans that were found stranded or severely entangled in fishing gear during the period 2004 through 2009. Approximately 57% of the bottlenose dolphins and 36% of the rough-toothed dolphins had significant hearing deficits with a reduction in sensitivity equivalent to severe (70-90 dB) or profound (>90 dB) hearing loss in humans. The only stranded short-finned pilot whale examined had profound hearing loss. No impairments were detected in seven Risso's dolphins from three different stranding events, two pygmy killer whales, one Atlantic spotted dolphin, one spinner dolphin, or a juvenile Gervais' beaked whale. Hearing impairment could play a significant role in some cetacean stranding events, and the hearing of all cetaceans in rehabilitation should be tested.     

Harassment and killing of porpoises (“phocoenacide”) by fish-eating Southern Resident killer whales (Orcinus orca)

Endangered Southern Resident killer whales (Orcinus orca) are fish-eaters that preferentially prey on adult Chinook salmon (Oncorhynchus tshawytscha). Despite being salmon specialists, individuals from all three killer whale pods (J, K, L) have been observed harassing and killing porpoises (family Phocoenidae) without consuming them. Retrospectively, we identified and analyzed 78 episodes of Southern Resident killer whales harassing porpoises between 1962 and 2020, of which 28 resulted in the porpoise's death (“phocoenacide”). Fifty-six episodes involved harbor porpoise (Phocoena phocoena), 13 involved Dall's porpoise (Phocoenoides dalli), and the porpoise species was unreported for nine episodes. Southern Resident killer whales often targeted young porpoises that were similar in size to adult Chinook salmon. Both sexes participated in porpoise harassment. Juveniles engaged in the behavior the most; however, their rates of engagement were not found to differ significantly from most other age classes. The behavior was passed through generations and social groupings, as it was first observed in L pod and spread to the other two pods. Killer whales are highly complex animals known to exhibit social learning and cultural transmission of learned behaviors, but the reason(s) for this behavior is unknown. Hypotheses include the social and developmental benefits of play, hunting practice, or displaced epimeletic behavior.     

Odontocete occurrence in relation to changes in oceanography at a remote equatorial Pacific seamount

Seamounts are considered hot spots of biodiversity and can aggregate pelagic predators and their prey. Passive acoustic monitoring was conducted over 3 mo in 2012 to document the occurrence of odontocetes near a seamount chain in the central equatorial Pacific in relation to oceanographic changes over time. Beaked whale echolocation signals were most frequently encountered. The main beaked whale signal was an unknown type, BW38, which resembled signals produced by Blainville's beaked whales. It had high occurrence during high sea surface temperature and low sea surface salinity. Cuvier's beaked whales were the second most detected. They had an opposite pattern and were encountered more often when sea surface temperature was low and net primary productivity was high. Risso's dolphins and short‐finned pilot whales had high acoustic densities, and echolocated predominantly at night. Risso's dolphins occurred more often during low sea surface height deviation. False killer whales were less frequently detected and mostly occurred during the day. Sperm whale detections were fewer than expected and associated with high chlorophyll a. Short duration Kogiidae encounters occurred on average every third day. These types of long‐term site studies are an informative tool to comparatively assess species composition, relative abundance, and relationship to oceanographic changes.     

SEASONAL PATTERNS IN THE ABUNDANCE AND DISTRIBUTION OF CALIFORNIA CETACEANS, 1991–1992

This study presents a detailed seasonal comparison of the abundance and distribution of cetaceans within 100-150 nmi (185-278 km) of the California coast during 1991 and 1992. The results of a shipboard line-transect survey conducted in July-November 1991 ("summer") were compared to those from aerial line-transect surveys conducted in March-April 1991 and February-April 1992 ("winter"). Using a confidence-interval-based bootstrap procedure, abundance estimates for six of the eleven species included in the comparison exhibited significant (α= 0.05) differences between the winter and summer surveys. Pacific white-sided dolphins ( Lagenorhynchus obliquidens ), Risso's dolphins ( Grampus griseus ), common dolphins ( Delphinus spp.), and northern right whale dolphins ( Lissodelphis borealis ) were significantly more abundant in winter. The abundance of blue whales ( Balaenoptera musculuss ) and gray whales ( Eschrichtius robustus ) reflected well-documented migratory patterns. Fin whales ( B. physalus ) were significantly more abundant during summer. No significant differences in seasonal abundance were identified for Dall's porpoises ( Phocoenoides dalli ), bottlenose dolphins ( Tursiops truncatus ), killer whales ( Orcinus orca ), sperm whales ( Physeter macrocephalus ), or humpback whales ( Megaptera novaeangliae ). Significant north/south shifts in distribution were found for Dall's porpoises, common dolphins, and Pacific white-sided dolphins, and significant inshore/offshore differences were identified for northern right whale dolphins and humpback whales.     

Age structure of strandings and growth of Lahille's bottlenose dolphin (Tursiops truncatus gephyreus)

This study describes the age structure and sex-specific growth patterns of Lahille's bottlenose dolphins (Tursiops truncatus gephyreus), a subspecies endemic to the Southwest Atlantic Ocean (SWAO). The ages of 120 animals collected in southern Brazil between 1976 and 2017 were determined. We found high frequencies of young animals, mostly males, with no sex bias among adults. A temporal change in the age structure of strandings was observed, and we conclude it is a consequence of increased bycatch rates of young dolphins after 2000. The oldest male and female were 27 and 44 years old, respectively, suggesting that females live longer than males in southern Brazil. Growth curve analysis using Gompertz and Laird-Gompertz growth models estimated asymptotic lengths of 316.5 cm for females and 351.6 cm for males, (reached around 13 and 18 years old, respectively), supporting sexual size dimorphism for the subspecies. A second growth pulse was identified for males. Our work highlights the benefit of long-term studies and contributes valuable information toward understanding the life history of Lahille's bottlenose dolphin. It will serve as baseline for future studies that seek to understand mortality patterns of bottlenose dolphins elsewhere and the effects of anthropogenic actions on the survival of these animals.     

Social evolution in toothed whales

Two contrasting results emerge from comparisons of the social systems of several odontocetes with terrestrial mammals. Researchers have identified remarkable convergence in prominent features of the social systems of odontocetes such as the sperm whale and bottlenose dolphin with a few well-known terrestrial mammals such as the elephant and chimpanzee. In contrast, studies on killer whales and Baird's beaked whale reveal novel social solutions to aquatic living. The combination of convergent and novel features in odontocete social systems promise a more general understanding of the ecological determinants of social systems in both terrestrial and aquatic habitats, as well as the relationship between relative brain size and social evolution.     

Living in the fast lane: rapid development of the locomotor muscle in immature harbor porpoises (Phocoena phocoena)

Cetaceans (dolphins and whales) are born into the aquatic environment and are immediately challenged by the demands of hypoxia and exercise. This should promote rapid development of the muscle biochemistry that supports diving, but previous research on two odontocete (toothed whales and dolphins) species showed protracted postnatal development for myoglobin content and buffering capacity. A minimum of 1 and 1.5 years were required for Fraser’s (Lagenodelphis hosei) and bottlenose (Tursiops truncatus) dolphins to obtain mature myoglobin contents, respectively; this corresponded to their lengthy 2 and 2.5-year calving intervals (a proxy for the dependency period of cetacean calves). To further examine the correlation between the durations for muscle maturation and maternal dependency, we measured myoglobin content and buffering capacity in the main locomotor muscle (longissimus dorsi) of harbor porpoises (Phocoena phocoena), a species with a comparatively short calving interval (1.5 years). We found that at birth, porpoises had 51 and 69 % of adult levels for myoglobin and buffering capacity, respectively, demonstrating greater muscle maturity at birth than that found previously for neonatal bottlenose dolphins (10 and 65 %, respectively). Porpoises achieved adult levels for myoglobin and buffering capacity by 9–10 months and 2–3 years postpartum, respectively. This muscle maturation occurred at an earlier age than that found previously for the dolphin species. These results support the observation that variability in the duration for muscular development is associated with disparate life history patterns across odontocetes, suggesting that the pace of muscle maturation is not solely influenced by exposure to hypoxia and exercise. Though the mechanism that drives this variability remains unknown, nonetheless, these results highlight the importance of documenting the species-specific physiological development that limits diving capabilities and ultimately defines habitat utilization patterns across age classes.     

Auditory temporal resolution and evoked responses to pulsed sounds for the Yangtze finless porpoises (<Emphasis Type="Italic">Neophocaena phocaenoides asiaeorientalis</Emphasis>)

Temporal cues are important for some forms of auditory processing, such as echolocation. Among odontocetes (toothed whales, dolphins, and porpoises), it has been suggested that porpoises may have temporal processing abilities which differ from other odontocetes because of their relatively narrow auditory filters and longer duration echolocation signals. This study examined auditory temporal resolution in two Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis) using auditory evoked potentials (AEPs) to measure: (a) rate following responses and modulation rate transfer function for 100 kHz centered pulse sounds and (b) hearing thresholds and response amplitudes generated by individual pulses of different durations. The animals followed pulses well at modulation rates up to 1,250 Hz, after which response amplitudes declined until extinguished beyond 2,500 Hz. The subjects had significantly better hearing thresholds for longer, narrower-band pulses similar to porpoise echolocation signals compared to brief, broadband sounds resembling dolphin clicks. Results indicate that the Yangtze finless porpoise follows individual acoustic signals at rates similar to other odontocetes tested. Relatively good sensitivity for longer duration, narrow-band signals suggests that finless porpoise hearing is well suited to detect their unique echolocation signals.     

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow,Acoustically Complex Habitat

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPa_pp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPa_pp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.     

Historical and current records of aquarium cetaceans in China

The number of cetaceans housed in aquariums in China is increasing. Detailed information on the historical and current population status has not been reported, despite its importance for successful breeding and population management. Questionnaires were conducted between December 2006 and May 2009, and the information was used to construct studbooks. Our survey showed that 10 species had been introduced to aquariums since 1978, including 26 (with 15 in the current population) finless porpoises (Neophocaena phocaenoides), 5 (5) false killer whales (Pseudorca crassidens), 94 (80) common bottlenose dolphins (Tursiops truncatus), 48 (30) Indo-Pacific bottlenose dolphins (Tursiops aduncus), 36 (32) beluga whales (Delphinapterus leucas), 10 (10) pantropical spotted dolphins (Stenella attenuata), 8 (8) Risso's dolphins (Grampus griseus), 2 (2) short-finned pilot whales (Globicephala macrorhynchus), 2 (2) Pacific white-sided dolphins (Lagenorhynchus obliquidens), and 5 (0) baiji dolphins (Lipotes vexillifer). The number of cetaceans has increased markedly in the past 32 years, especially since 1995. Currently, 184 individuals are under human care throughout China, a number larger than any other country with an International Species Information System membership. In addition, the Annual Survival Rates of bottlenose dolphins (0.959) and beluga whales (0.968) were found higher than those reported previously (0.93-0.951 and 0.94-0.954, respectively).     

Life history constrains biochemical development in the highly specialized odontocete echolocation system

The vertebrate head has undergone enormous modification from the features borne by early ancestors. The growth of skull bones has been well studied in many species, yet little is known about corresponding soft tissue development. Among mammals, some of the most unusual examples of cranial evolution exist in the toothed whales (odontocetes). Specialized fat bodies in toothed whale heads play important roles in sound transmission and reception. These fat bodies contain unique endogenous lipids, with favourable acoustic properties, arranged in highly organized, three-dimensional patterns. We link variation in developmental rates of acoustic fats with life-history strategy, using bottlenose dolphins and harbour porpoises. Porpoise acoustic fats attain adult configurations earlier (less than 1 year) and at a faster pace than dolphins. The accelerated lipid accumulation in porpoises reflects the earlier need for fully functional echolocation systems. Dolphins enjoy 3-6 years of maternal care; porpoises must achieve total independence by approximately nine months. Further, a stereotypic 'blueprint' for the spatial distribution of lipids is established prior to birth, demonstrating the highly conserved nature of the intricate biochemical arrangement in acoustic tissues. This system illustrates an unusual case of soft tissue development being constrained by life history, rather than the more commonly observed mechanistic or phyletic constraints.     

Ecotype and geographical variation in carbon and nitrogen stable isotope values in western North Pacific killer whales (Orcinus orca)

Killer whales are top predators in marine trophic chains, and therefore their feeding preferences can substantially affect the abundance of species on the lower trophic levels. Killer whales are known to feed on many different types of prey from small fish to large whales, but a given killer whale population usually focuses on a specific type of prey. Stable isotope analysis is widely used to study whale diets, because direct observations are often impossible. Killer whale feeding habits in the western North Pacific are poorly studied, and the large-scale stable isotope analysis provides a unique opportunity to gain insights into the trophic links of this top predator. In this study, we compare the δ¹³C and δ¹⁵N stable isotope values from killer whale skin samples obtained in different areas of the western North Pacific from fish-eating (R-type) and mammal-eating (T-type) killer whale ecotypes. The effect of ecotype was highly significant: both carbon and nitrogen stable isotope values were lower in R-type whales than in T-type whales. The geographical variation also affected killer whale stable isotope values due to both the differences in killer whale diet and the variation in baseline stable isotope values across the study areas.