Body and self in dolphins.

In keeping with recent views of consciousness of self as represented in the body in action, empirical studies are reviewed that demonstrate a bottlenose dolphin's (Tursiops truncatus) conscious awareness of its own body and body parts, implying a representational "body image" system. Additional work reviewed demonstrates an advanced capability of dolphins for motor imitation of self-produced behaviors and of behaviors of others, including imitation of human actions, supporting hypotheses that dolphins have a sense of agency and ownership of their actions and may implicitly attribute those levels of self-awareness to others. Possibly, a mirror-neuron system, or its functional equivalent to that described in monkeys and humans, may mediate both self-awareness and awareness of others.

Sound production by singing humpback whales.

Sounds from humpback whale songs were analyzed to evaluate possible mechanisms of sound production. Song sounds fell along a continuum with trains of discrete pulses at one end and continuous tonal signals at the other. This graded vocal repertoire is comparable to that seen in false killer whales [Murray et al. (1998). J. Acoust. Soc. Am. 104, 1679-1688] and human singers, indicating that all three species generate sounds by varying the tension of pneumatically driven, vibrating membranes. Patterns in the spectral content of sounds and in nonlinear sound features show that resonating air chambers may also contribute to humpback whale sound production. Collectively, these findings suggest that categorizing individual units within songs into discrete types may obscure how singers modulate song features and illustrate how production-based characterizations of vocalizations can provide new insights into how humpback whales sing.

The dolphin's (Tursiops truncatus) understanding of human gazing and pointing: knowing what and where.

The authors tested whether the understanding by dolphins (Tursiops truncatus) of human pointing and head-gazing cues extends to knowing the identity of an indicated object as well as its location. In Experiment 1, the dolphins Phoenix and Akeakamai processed the identity of a cued object (of 2 that were present), as shown by their success in selecting a matching object from among 2 alternatives remotely located. Phoenix was errorless on first trials in this task. In Experiment 2, Phoenix reliably responded to a cued object in alternate ways, either by matching it or by acting directly on it, with each type of response signaled by a distinct gestural command given after the indicative cue. She never confused matching and acting. In Experiment 3, Akeakamai was able to process the geometry of pointing cues (but not head-gazing cues), as revealed by her errorless responses to either a proximal or distal object simultaneously present, when each object was indicated only by the angle at which the informant pointed. The overall results establish that these dolphins could identify, through indicative cues alone, what a human is attending to as well as where.

The multiple functions of male song within the humpback whale (Megaptera novaeangliae) mating system: review, evaluation, and synthesis.

Humpback whales (Megaptera novaeangliae) are seasonal breeders, annually migrating from high-latitude summer feeding grounds to low-latitude winter breeding grounds. The social matrix on the winter grounds is a loose network of interacting individuals and groups and notably includes lone males that produce long bouts of complex song that collectively yield an asynchronous chorus. Occasionally, a male will sing while accompanying other whales. Despite a wealth of knowledge about the social matrix, the full characterization of the mating system remains unresolved, without any firm consensus, as does the function of song within that system. Here, I consider and critically analyse three proposed functions of song that have received the most attention in the literature: female attraction to individual singers, determining or facilitating male-male interactions, and attracting females to a male aggregation within the context of a lekking system. Female attraction suggests that humpback song is an advertisement and invitation to females, but field observations and song playback studies reveal that female visits to individual singers are virtually absent. Other observations suggest instead that females might convey their presence to singers (or to other males) through the percussive sounds of flipper or tail slapping or possibly through vocalizations. There is some evidence for male-male interactions, both dominance and affiliative: visits to singers are almost always other lone males not singing at that time. The joiner may be seeking a coalition with the singer to engage cooperatively in attempts to obtain females, or may be seeking to disrupt the song or to affirm his dominance. Some observations support one or the other intent. However, other observations, in part based on the brevity of most pairings, suggest that the joiner is prospecting, seeking to determine whether the singer is accompanying a female, and if not soon departs. In the lekking hypothesis, the aggregation of vocalizing males on a winter ground and the visits there by non-maternal females apparently for mating meet the fundamental definition of a lekking system and its role though communal display in attracting females to the aggregation, although not to an individual singer. Communal singing is viewed as a form of by-product mutualism in which individuals benefit one another as incidental consequences of their own selfish actions. Possibly, communal singing may also act to stimulate female receptivity. Thus, there are both limitations and merit in all three proposals. Full consideration of song as serving multiple functions is therefore necessary to understand its role in the mating system and the forces acting on the evolution of song. I suggest that song may be the prime vector recruiting colonists to new winter grounds pioneered by vagrant males as population pressures increase or as former winter grounds become unavailable or undesirable, with such instances documented relatively recently. Speculatively, song may have evolved historically as an aggregating call during the dynamic ocean conditions and resulting habitat uncertainties in the late Miocene-early Pliocene epochs when Megaptera began to proliferate. Early song may have been comprised of simpler precursor sounds that through natural selection and ritualization evolved into complex song.

Bottlenosed dolphins (Tursiops truncatus) comprehend the referent of both static and dynamic human gazing and pointing in an object-choice task.

The authors tested 2 bottlenosed dolphins (Tursiops truncatus) for their understanding of human-directed gazing or pointing in a 2-alternative object-choice task. A dolphin watched a human informant either gazing at or pointing toward 1 of 2 laterally placed objects and was required to perform a previously indicated action to that object. Both static and dynamic gaze, as well as static and dynamic direct points and cross-body points, yielded errorless or nearly errorless performance. Gaze with the informant's torso obscured (only the head was shown) produced no performance decrement, but gaze with eyes only resulted in chance performance. The results revealed spontaneous understanding of human gaze accomplished through head orientation, with or without the human informant's eyes obscured, and demonstrated that gaze-directed cues were as effective as point-directed cues in the object-choice task.

Recurring patterns in the songs of humpback whales (Megaptera novaeangliae).

Humpback whales, unlike most mammalian species, learn new songs as adults. Populations of singers progressively and collectively change the sounds and patterns within their songs throughout their lives and across generations. In this study, humpback whale songs recorded in Hawaii from 1985 to 1995 were analyzed using self-organizing maps (SOMs) to classify the sounds within songs, and to identify sound patterns that were present across multiple years. These analyses supported the hypothesis that recurring, persistent patterns exist within whale songs, and that these patterns are defined at least in part by acoustic relationships between adjacent sounds within songs. Sound classification based on acoustic differences between adjacent sounds yielded patterns within songs that were more consistent from year to year than classifications based on the properties of single sounds. Maintenance of fixed ratios of acoustic modulation across sounds, despite large variations in individual sounds, suggests intrinsic constraints on how sounds change within songs. Such acoustically invariant cues may enable whales to recognize and assess variations in songs despite propagation-related distortion of individual sounds and yearly changes in songs.

A claim in search of evidence: reply to Manger's thermogenesis hypothesis of cetacean brain structure.

In a recent publication in Biological Reviews, Manger (2006) made the controversial claim that the large brains of cetaceans evolved to generate heat during oceanic cooling in the Oligocene epoch and not, as is the currently accepted view, as a basis for an increase in cognitive or information-processing capabilities in response to ecological or social pressures. Manger further argued that dolphins and other cetaceans are considerably less intelligent than generally thought. In this review we challenge Manger's arguments and provide abundant evidence that modern cetacean brains are large in order to support complex cognitive abilities driven by social and ecological forces.

Dolphin (Tursiops truncatus) echoic angular discrimination: effects of object separation and complexity.

A bottlenose dolphin was tested on its ability to echoically discriminate horizontal angular differences between arrays of vertically oriented air-filled PVC rods. The blindfolded dolphin was required to station in a submerged hoop 2 radial m from the stimuli and indicate if an array with two rods (S+) was to the right or the left of a single rod (S-). The angular separation between the two rods (thetaw) was held constant within each experiment while the angle between the S+ and the S-stimuli (thetab) varied to produce angular differences (deltatheta= thetab-thetaw) ranging from 0.25 to 4 degrees. In experiment I, thetaw was maintained at 2 degrees and in experiment II, thetaw was maintained at 4 degrees. Resulting 75% correct thresholds (method of constant stimuli) were 1.5 and 0.7 degrees, respectively. The two main findings of this study are: (1) decreasing the number of targets does not aid in localization, and (2) increasing the space between the rods enhances localization. Taken as a whole, the experiments suggest dolphins have a well-developed ability to resolve spatial information through sonar.

Acoustic properties of humpback whale songs.

A vertical array of five hydrophones was used to measure the acoustic field in the vertical plane of singing humpback whales. Once a singer was located, two swimmers with snorkel gear were deployed to determine the orientation of the whale and position the boat so that the array could be deployed in front of the whale at a minimum standoff distance of at least 10 m. The spacing of the hydrophones was 7 m with the deepest hydrophone deployed at a depth of 35 m. An eight-channel TASCAM recorder with a bandwidth of 24 kHz was used to record the hydrophone signals. The location (distance and depth) of the singer was determined by computing the time of arrival differences between the hydrophone signals. The maximum source level varied between individual units in a song, with values between 151 and 173 dB re 1 microPa. One of the purposes of this study was to estimate potential sound exposure of nearby conspecifics. The acoustic field determined by considering the relative intensity of higher frequency harmonics in the signals indicated that the sounds are projected in the horizontal direction despite the singer being canted head downward anywhere from about 25 degrees to 90 degrees. High-frequency harmonics extended beyond 24 kHz, suggesting that humpback whales may have an upper frequency limit of hearing as high as 24 kHz.

Song copying by humpback whales: themes and variations.

Male humpback whales (Megaptera novaeangliae) produce long, structured sequences of sound underwater, commonly called "songs." Humpbacks progressively modify their songs over time in ways that suggest that individuals are copying song elements that they hear being used by other singers. Little is known about the factors that determine how whales learn from their auditory experiences. Song learning in birds is better understood and appears to be constrained by stable core attributes such as species-specific sound repertoires and song syntax. To clarify whether similar constraints exist for song learning by humpbacks, we analyzed changes over 14 years in the sounds used by humpback whales singing in Hawaiian waters. We found that although the properties of individual sounds within songs are quite variable over time, the overall distribution of certain acoustic features within the repertoire appears to be stable. In particular, our findings suggest that species-specific constraints on temporal features of song sounds determine song form, whereas spectral variability allows whales to flexibly adapt song elements.

The object behind the echo: dolphins (Tursiops truncatus) perceive object shape globally through echolocation.

Two experiments tested a bottlenosed dolphin's ability to match objects across echolocation and vision. Matching was tested from echolocation sample to visual alternatives (E-V) and from visual sample to echolocation alternatives (V-E). In Experiment 1, the dolphin chose a match from among three-alternative objects that differed in overall (global) shape, but shared several 'local' features with the sample. The dolphin conducted a right-to-left serial nonexhaustive search among the alternatives, stopping when a match was encountered. It matched correctly on 93% of V-E trials and on 99% of E-V trials with completely novel combinations of objects despite the presence of many overlapping features. In Experiment 2, a fourth alternative was added in the form of a paddle that the dolphin could press if it decided that none of the three-alternatives matched the sample. When a match was present, the dolphin selected it on 94% of V-E trials and 95% of E-V trials. When a match was absent, the dolphin pressed the paddle on 74% and 76%, respectively, of V-E and E-V trials. The approximate 25% error rate, which consisted of a choice of one of the three non-matching alternatives in lieu of the paddle press, increased from right to center to left alternative object, reflecting successively later times in the dolphin's search path. A weakening in memory for the sample seemed the most likely cause of this error pattern. Overall, the results gave strong support to the hypothesis that the echolocating dolphin represents an object by its global appearance rather than by local features.