Why do dolphins jump? Interpreting the behavioural repertoire of bottlenose dolphins (Tursiops sp.) in Doubtful Sound,New Zealand

Only a limited number of studies have tried to determine the purpose of surface behavioural events performed by dolphins. To date only one study has attempted to aggregate the behavioural events observed in a population in contextual groups using co-occurrence as the grouping factor. In the present study, I tried to characterise the behavioural repertoire of a bottlenose dolphin population (Tursiops sp.) present in Doubtful Sound, New Zealand. I first looked at the relationship between events performed by individuals depending on the behavioural state of their schools. I then assessed the likelihood for events to co-occur. Four main behavioural categories (orientation, travel, social displays and fights) emerged from this analysis. Aerial events (jumps) did not fall into one category, showing that different aerial behaviours play different roles. Moreover, it appears that dolphins used side-flopping and upside-down lobtailing to communicate motivation. Side-flops occurred when the focal schools finished a behavioural bout and started to travel, while upside-down lobtails occurred when the focal schools instigated a behavioural bout after travelling. This non-vocal communication can take place over a few meters to hundreds of meters. Having signals that are effective over very short ranges avoids unwanted signalling to prey, predators or conspecifics.     

Distribution and abundance of Indo‐Pacific humpback dolphins in Leizhou Bay,China

Abstract

The Indo‐Pacific humpback dolphin (Sousa chinensis) is under severe threat in Chinese waters. Only four small populations of this dolphin are known to survive in the East China Sea and South China Sea. We conducted exploratory surveys from June to September 2005, and found a population of humpback dolphins in Leizhou Bay. Thirty‐eight individuals were identified and catalogued, of which 26 were photographed and re‐identified on more than one occasion, and nine were sighted on five or more occasions each. Our preliminary estimate of the size of the Leizhou population is about 237 individuals, second only to the Pearl River estuary population. We suggest that Leizhou Bay has the potential to serve as a “humpback dolphin sanctuary” in Chinese waters.     

Genetic differentiation and intraspecific structure of Eastern Tropical Pacific spotted dolphins,Stenella attenuata,revealed by DNA analyses

Mitochondrial DNA (mtDNA) control region sequences and microsatellite loci length polymorphisms were used to investigate genetic differentiation in spotted dolphins (Stenella attenuata) in the Eastern Tropical Pacific and to examine the intraspecific structure of the coastal subspecies (Stenella attenuata graffmani). One-hundred and thirty-five animals from several coastal areas and 90 offshore animals were sequenced for 455 bp of the mitochondrial control region, resulting in 112 mtDNA haplotypes. Phylogenetic analyses and the existence of shared haplotypes between the two subspecies suggest recent and/or current gene flow. Analyses using χ², F _ST (based on haplotype frequencies) and Φ_ST values (based on frequencies and genetic distances between haplotypes) yielded statistically significant separation (randomized permutation values P<0.05) among four different coastal populations and between all but one of these and the offshore subspecies (overall F _ST=0.0691). Ninety-one coastal animals from these four geographic populations and 50 offshore animals were genotyped for seven nuclear microsatellite loci. Analysis using F _ST values (based on allelic frequencies) yielded statistically significant separation between most coastal populations and offshore animals, although no coastal populations were distinguished. These results argue for the existence of some genetic isolation between offshore and inshore populations and among some inshore populations, suggesting that these should be treated as separate units for management purposes.     

Spatial distribution and aggregation of Xenobalanus globicipitis on the flukes of striped dolphins,Stenella coeruleoalba: An indicator of host hydrodynamics?

We analyzed patterns of aggregation and spatial distribution of the epizoic barnacle Xenobalanus globicipitis on the flukes of Mediterranean striped dolphins, Stenella coeruleoalba, assessing its potential use as an indicator of the host's hydrodynamics based on data from 55 dolphins. Barnacles occurred along the trailing edge with the cirri oriented towards the oncoming flow. Nearest neighbor analyses suggested that new recruits actively seek placement next to already settled barnacles, forming aggregations possibly to facilitate copulation. The probability of spanwise settlement strongly increased with fluke width (presumably enabling prolonged leading-edge vorticity), and with chordwise length where pressure is predicted to be positive. Consequently, clustering tended to increase nonlinearly towards the fluke notch. Furthermore, it was three times more likely for barnacles to occur on the dorsal vs. ventral side of flukes, at an average abundance ca. 12 times higher. This difference could result from a host's asymmetric oscillation of the tail, which would alter leading-edge vorticity, and/or an interaction between the wake produced by the dorsal fin and the flow associated with flukes. Both processes could primarily enhance cyprid contact and/or attachment on the dorsal side. This study offers a starting point for future comparison with other cetaceans.     

Communication in bottlenose dolphins: 50 years of signature whistle research

Bottlenose dolphins (Tursiops truncatus) produce individually distinctive signature whistles that broadcast the identity of the caller. Unlike voice cues that affect all calls of an animal, signature whistles are distinct whistle types carrying identity information in their frequency modulation pattern. Signature whistle development is influenced by vocal production learning. Animals use a whistle from their environment as a model, but modify it, and thus invent a novel signal. Dolphins also copy signature whistles of others, effectively addressing the whistle owner. This copying occurs at low rates and the resulting copies are recognizable as such by parameter variations in the copy. Captive dolphins can learn to associate novel whistles with objects and use these whistles to report on the presence or absence of the object. If applied to signature whistles, this ability would make the signature whistle a rare example of a learned referential signal in animals. Here, we review the history of signature whistle research, covering definitions, acoustic features, information content, contextual use, developmental aspects, and species comparisons with mammals and birds. We show how these signals stand out amongst recognition calls in animals and how they contribute to our understanding of complexity in animal communication.     

Can dolphins heal by ultrasound?

In recent years, dolphin-assisted therapy has become very popular and an increasing number of facilities offer therapy programs with dolphins worldwide. To this date, there are no studies concerning the behavior of dolphins during these therapies. As a result of speculations that the echolocation of dolphins may play an important role for the success of the therapy and the high publicity of this in the media, people pay much more for dolphin-assisted than for other animal-assisted therapy programs. Based on publications in medicine, we will show that ultrasound emitted by dolphins could have an effect on biological tissue under some circumstances; such as sufficient intensity, repeated application over several days or weeks and a certain application duration per session. We recorded 83 sessions at the “Dolphins Plus”, a fenced area with ocean water in the Florida Keys. Our observations demonstrate that only one out of five observed dolphins behave significantly differently towards patients compared to other humans and that the duration of the observed close contacts did not meet the requirements for common ultrasound therapies.     

Do dolphins know their own minds?

Knowledge of one's own states of mind is one of the varieties of self-knowledge. Do any nonhuman animals have the capacity for this variety of self-knowledge? The question is open to empirical inquiry, which is most often conducted with primate subjects. Research with a bottlenose dolphin gives some evidence for the capacity in a nonprimate taxon. I describe the research and evaluate the metacognitive interpretation of the dolphin's behaviour. The research exhibits some of the difficulties attached to the task of eliciting behaviour that both attracts a higher-order interpretation while also resisting deflationary, lower-order interpretations. Lloyd Morgan's Canon, which prohibits inflationary interpretations of animal behaviour, has influenced many animal psychologists. There is one defensible version of the Canon, the version that warns specifically against unnecessary intentional ascent. The Canon on this interpretation seems at first to tell against a metacognitive interpretation of the data collected in the dolphin study. However, the model of metacognition that is in play in the dolphin studies is a functional model, one that does not implicate intentional ascent. I explore some interpretations of the dolphin's behaviour as metacognitive, in this sense. While this species of metacognitive interpretation breaks the connection with the more familiar theory of mind research using animal subjects, the interpretation also points in an interesting way towards issues concerning consciousness in dolphins.     

Whistle variability of Guiana dolphins in South America: Latitudinal variation or acoustic adaptation?

A previous comparison of whistles using data sampled at 48 kHz suggested that certain frequency parameters vary along a latitudinal gradient. This geographical pattern may be biased because whistles sampled at higher frequencies could potentially have very different frequency contents. The current study compared the acoustic parameters of Guiana dolphin (Sotalia guianensis) whistles recorded at a higher sampling rate (96 kHz) and from groups occupying two never before sampled sites, Benevente Bay, Espírito Santo, Brazil, and Formosa Bay, Rio Grande do Norte, Brazil, with recordings of other populations in South America. By only considering data sampled at a rate of at least 96 kHz, we aimed to detect differences in whistles across locations. Contrary to previous findings, our analyses do not indicate any clear separation between northern and southern populations based on whistles, and do not corroborate the hypothesis of latitudinal acoustic variation in this species. The variation in Guiana dolphin whistle parameters found here appears to be influenced by latitude to some extent, but several other factors, including sampling method, environmental fluctuations, and social influence on vocal learning, may be confounding the detection of a geographic pattern in these whistle samples.     

Diet of common dolphins (Delphinus delphis) off the south-east coast of southern Africa: opportunism or specialization?

A total of 297 common dolphin (Delphinus delphis) stomachs was used for these diet analyses. All originated from common dolphins incidentally captured in Natal, where their occurrence is strongly associated with an annual, northward fish migration, the sardine run, along the east coast. Thirty-six fish and four cephalopod prey species were identified. Though 86.9% by weight of the diet was made up of only five prey species, common dolphins appear to feed opportunistically, their diet reflecting local prey abundance and availability. Prey were primarily small, easily-caught, pelagic shoaling species, with the main prey being South African pilchard (Sardinops ocellatus). Although pilchard dominated the diet between 1974 and 1992, there were marked annual and seasonal fluctuations in dietary composition, reflecting changes in prey availability and abundance. There were clear differences in the contribution of particular prey species to the diet of different sex and size groups, with strong evidence for resource partitioning between groups. The influence of the annual 'sardine run', and the fish predators associated with it, on the diet, feeding biology and distribution of common dolphins is discussed.     

'Eavesdropping' in wild rough-toothed dolphins (Steno bredanensis)?

Several authors suggest that dolphins use information obtained by eavesdropping on echoes from sonar signals of conspecifics, but there is little evidence that this strategy is used by dolphins in the wild. Travelling rough-toothed dolphins (Steno bredanensis) either exhibit asynchronous movements or an extremely synchronized swimming behaviour in tight formations, which we expect to facilitate eavesdropping. Therefore, we determined, whether either one or more dolphins were echolocating in subgroups that were travelling with asynchronous and synchronized movements. Since, the number of recording sequences in which more than one animal produced sonar signals was significantly lower during synchronized travel, we conclude that the other members of a subgroup might get information on targets ahead by eavesdropping. Synchronized swimming in tight formations might be an energetic adaptation for travelling in a pelagic dolphin species that facilitates eavesdropping.     

What can dolphins tell us about primate evolution?

Fifty-five million years ago, a furry, hoofed mammal about the size of a dog ventured into the shallow brackish remnant of the Tethys Sea and set its descendants on a path that would lead to their complete abandonment of the land. These early ancestors of cetaceans (dolphins, porpoises, and whales) thereafter set on an evolutionary course that is arguably the most unusual of any mammal that ever lived. Primates and cetaceans, because of their adaptation to exclusively different physical environments, have had essentially nothing to do with each other throughout their evolution as distinct orders. In fact, the closest phylogenetic relatives of cetaceans are even-toed ungulates.     

To feed or not to feed? Bioenergetic impacts of fear‐driven behaviors in lactating dolphins

In mammals, lactation can be the most energetically expensive part of the reproductive cycle. Thus, when energy needs are compromised due to predation risk, environmental disturbance, or resource scarcity, future reproductive success can be impacted. In marine and terrestrial environments, foraging behavior is inextricably linked to predation risk. But quantification of foraging energetics for lactating animals under predation risk is less understood. In this study, we used a spatially explicit individual‐based model to study how changes in physiology (lactating or not) and the environment (predation risk) affect optimal behavior in dolphins. Specifically, we predicted that an adult dolphin without calf would incur lower relative energetic costs compared to a lactating dolphin with calf regardless of predation risk severity, antipredator behavior, or prey quality consumed. Under this state‐dependent analysis of risk approach, we found predation risk to be a stronger driver in affecting total energetic costs (foraging plus locomotor costs) than food quality for both dolphin types. Further, contrary to our hypothesis, after accounting for raised energy demands, a lactating dolphin with calf does not necessarily have higher relative‐to‐baseline costs than a dolphin without calf. Our results indicate that both a lactating (with calf) and non‐lactating dolphin incur lowered energetic costs under a risk‐averse behavioral scheme, but consequently suffer from lost foraging calories. A lactating dolphin with calf could be particularly worse off in lost foraging calories under elevated predation risk, heightened vigilance, and increased hiding time relative to an adult dolphin without calf. Further, hiding time in refuge could be more consequential than detection distance for both dolphin types in estimated costs and losses incurred. In conclusion, our study found that reproductive status is an important consideration in analyzing risk effects in mammals, especially in animals with lengthy lactation periods and those exposed to both biological and nonbiological stressors.     

Specialized sound reception in dolphins—a hint for the function of the dorsal cochlear nucleus in mammals

Dolphins have developed a specialized system for sound reception. Their pinnae are reduced and shifted under the skin, and sound is transduced via the lower jaw to the auditory ossicles. In line with their auditory functions (notably echolocation), most auditory nuclei in the brain stem of dolphins are extremely well‐developed. In contrast, the dorsal cochlear nucleus is drastically reduced. In the La Plata dolphin the volume of the dorsal cochlear nucleus is 4 mm³, while that of the ventral cochlear nucleus is 98 mm³. The reduction in size of the dorsal cochlear nucleus is also seen in baleen whales, which do not echolocate and are, in contrast to dolphins, tuned to deep frequencies. However, dolphins and baleen whales have the reduction of the outer ear in common. In humans, cats, bats and seals, there is a correlation between the development of pinnae and the dorsal cochlear nucleus. So, we conclude that the function of the dorsal cochlear nucleus in mammals could be to eliminate “auditory artifacts”; arising from the operation of the pinnae.     

Staying close to home? Genetic differentiation of rough-toothed dolphins near oceanic islands in the central Pacific Ocean

Rough-toothed dolphins have a worldwide tropical and subtropical distribution, yet little is known about the population structure and social organization of this typically open-ocean species. Although it has been assumed that pelagic dolphins range widely due to the lack of apparent barriers and unpredictable prey distribution, recent evidence suggests rough-toothed dolphins exhibit fidelity to some oceanic islands. Using the most comprehensively extensive dataset for this species to date, we assess the isolation and interchange of rough-toothed dolphins at the regional and oceanic scale within the central Pacific Ocean. Using mtDNA and microsatellite genotyping (nDNA), we analyzed samples of insular communities from the main Hawaiian (Kaua‘i n = 93, O‘ahu n = 9, Hawai‘i n = 57), French Polynesian (n = 70) and Samoan (n = 16) archipelagos, and pelagic samples off the Northwestern Hawaiian Islands (n = 18). An overall AMOVA indicated strong genetic differentiation among islands (mtDNA F_ST = 0.265; p < 0.001; nDNA F_ST = 0.038; p < 0.001), as well as among archipelagos (mtDNA F_ST = 0.299; p < 0.001; nDNA F_ST = 0.055; p < 0.001). Shared haplotypes (n = 4) between the archipelagos may be a product of a relatively recent divergence and/or periodic exchange from poorly understood pelagic populations. Analyses using STRUCTURE and GENELAND identified four separate management units among archipelagos and within the Hawaiian Islands. These results confirm the presence of multiple insular populations within the Pacific and island-specific genetic isolation among populations attached to islands in each archipelago. Insular populations seem most prevalent where oceanographic conditions indicate high local productivity or a discontinuity with surrounding oligotrophic areas. Our findings have important implications for a little studied species that faces increasing anthropogenic threats around oceanic islands.     

The bottlenose dolphin Tursiops truncatus foraging around a fish farm：Effects of prey abundance on dolphins＇ behavior

The extent to which prey abundance influences both bottlenose dolphin foraging behavior and group size in the presence of human activities has not previously been studied.The primary aim of this study was to identify and quantify how wild bottlenose dolphins respond,individually and as groups,to the relative abundance of prey around a fish farm.Detailed views of dolphins＇ behavior were obtained by focal following individual animals whilst simultaneously collecting surface and underwater behavioral data.A total of 2150 dive intervals were analyzed,corresponding to 342 focal samples,lasting over 34 hours.Bottlenose dolphins remained submerged for a mean duration of 46.4 seconds and a maximum of 249 seconds.This study provides the first quantified data on bottlenose dolphin diving behavior in a marine fin-fish farm area.This study＇s results indicate that within a fish farm area used intensively by bottlenose dolphins for feeding,dolphins did not modify dive duration.Additionally,underwater observations confirmed that dolphins find it easier to exploit a concentrated food source and it appears that hunting tactic and not group size plays an important role during feeding activities.Thus,bottlenose dolphins appear capable of modifying their hunting tactics according to the abundance of prey.When top predators display behavioral responses to activities not directed at them,the task of studying all possible effects of human activities can become even more challenging.     

Can habitat characteristics shape vertebral morphology in dolphins? An example of two phylogenetically related species from southern South America

Fast swimming pelagic cetacean species have osteological characteristics that promote a more stable spine in comparison to that of coastal species. The Peale's dolphin (Lagenorhynchus australis) and the hourglass dolphin (Lagenorhynchus cruciger) have a close phylogenetic relationship and are found in coastal and pelagic waters in the Southern Hemisphere, respectively. The aim of this work was to study the relationship between the vertebral column's morphology and its flexibility, across these species of contrasting habitats. Vertebral counts and multiple measurements of each vertebra were used to infer intervertebral flexibility. Bivariate plots and discriminant multivariate analyses were employed to compare each functional region along the vertebral column. Both species displayed a regionalization of the column into three stable regions and two flexible areas, which statistically differ in the proportion of the skeleton occupied in each species. While the Peale's dolphin has rounder vertebrae, associated with higher flexibility, the hourglass dolphin has disk‐shaped vertebrae and strongly inclined processes related to high stability. Although the species are closely related phylogenetically, vertebral morphology is influenced by a diverse set of ecological and behavioral factors, reflecting a high degree of vertebral plasticity within the genus.     

Genetic differentiation and limited gene flow among fragmented populations of New Zealand endemic Hector’s and Maui’s dolphins

Gene flow among small fragmented populations is critical for maintaining genetic diversity, and therefore the evolutionary potential of a species. Concern for two New Zealand endemic subspecies, the Hector’s (Cephalorhynchus hectori hectori) and Maui’s (C. h. maui) dolphins, arises from their low abundance, slow rate of reproduction, and susceptibility to fisheries-related mortality. Our work examined genetic differentiation and migration between the subspecies and among regional and local Hector’s dolphin populations using mitochondrial (mt) DNA and microsatellite genotypes from 438 samples. Results confirmed earlier reports of a single unique mtDNA control region haplotype fixed in the Maui’s dolphin, and provided new evidence of reproductive isolation from Hector’s dolphins (9-locus microsatellite F _ST?=?0.167, P?<?0.001). Independent evolutionary trajectories were also supported for Hector’s dolphin populations of the East Coast, West Coast, Te Waewae Bay and Toetoe Bay. Low asymmetrical migration rates were found among several Hector’s dolphin populations and assignment tests identified five Hector’s dolphins likely to have a migrant father from another regional population. There appears to be sufficient step-wise gene flow to maintain genetic diversity within the East and West Coasts; however, the two local South Coast populations exhibited a high degree of differentiation given their close proximity (~100?km). To maintain the evolutionary potential and long-term survival of both subspecies, genetic monitoring and conservation management must focus on maintaining corridors to preserve gene flow and prevent further population fragmentation and loss of genetic diversity, in addition to maintaining local population abundances.     

Integrating multiple lines of evidence to better understand the evolutionary divergence of humpback dolphins along their entire distribution range: a new dolphin species in Australian waters?

The conservation of humpback dolphins, distributed in coastal waters of the Indo‐West Pacific and eastern Atlantic Oceans, has been hindered by a lack of understanding about the number of species in the genus (Sousa) and their population structure. To address this issue, we present a combined analysis of genetic and morphologic data collected from beach‐cast, remote‐biopsied and museum specimens from throughout the known Sousa range. We extracted genetic sequence data from 235 samples from extant populations and explored the mitochondrial control region and four nuclear introns through phylogenetic, population‐level and population aggregation frameworks. In addition, 180 cranial specimens from the same geographical regions allowed comparisons of 24 morphological characters through multivariate analyses. The genetic and morphological data showed significant and concordant patterns of geographical segregation, which are typical for the kind of demographic isolation displayed by species units, across the Sousa genus distribution range. Based on our combined genetic and morphological analyses, there is convincing evidence for at least four species within the genus (S. teuszii in the Atlantic off West Africa, S. plumbea in the central and western Indian Ocean, S. chinensis in the eastern Indian and West Pacific Oceans, and a new as‐yet‐unnamed species off northern Australia).     

Checklist of the smaller families of Opomyzoidea,Anthomyzidae, Asteiidae,Aulacigastridae, Clusiidae,Odiniidae, Opomyzidae and Periscelididae (Diptera)?of?Finland

A species checklist is presented for Finland covering seven smaller families of Opomyzoidea: Anthomyzidae, Asteiidae, Aulacigastridae, Clusiidae, Odiniidae, Opomyzidae and Periscelididae (Diptera).