Joseph Soltis

African Elephant Vocal Communication II: Rumble Variation Reflects the Individual Identity and Emotional State of Callers

The most common vocalization of the African elephant, Loxodonta africana, is the rumble, but there is no consensus as to how many rumble subtypes exist. From the standpoint of social function, many types of rumble have been proposed. From a structural standpoint, however, few studies have examined detailed acoustic measurements of a large number of calls. We analysed 270 rumbles from six adult female African elephants housed at Disneys Animal Kingdom (Lake Buena Vista, Florida, U.S.A.). Subjects wore collars outfitted with microphones and radiotransmitters that allowed recording of vocalizations from identified individuals. Rumble vocalizations were digitized and both source and filter features were measured for each call. Behavioural and endocrine data were collected so that acoustical data could be placed into the context of ongoing social behaviour and reproductive state. Multidimensional scaling analysis revealed that, from a structural standpoint, rumbles from this captive setting could not be divided into distinct subtypes, but there was extensive acoustic variation across rumbles. Discriminant function analysis and MANOVA were employed to further explore this variation. First, acoustic characteristics varied according to the individual identity of the caller. Second, rumbles varied as a function of negative emotional arousal. When associating with dominant animals, subordinate females produced rumbles with lower cepstral coefficients, suggesting low tonality and unstable pitch in the voice, compared to rumbles produced outside of the presence of dominant animals. Rumbles as a whole did not cluster into distinct acoustic types, but structural variation in rumbles reflected the individual identity and emotional state of callers.

Stress and Emotion Classification using Jitter and Shimmer Features

In this paper, we evaluate the use of appended jitter and shimmer speech features for the classification of human speaking styles and of animal vocalization arousal levels. Jitter and shimmer features are extracted from the fundamental frequency contour and added to baseline spectral features, specifically Mel-frequency cepstral coefficients (MFCCs) for human speech and Greenwood function cepstral coefficients (GFCCs) for animal vocalizations. Hidden Markov models (HMMs) with Gaussian mixture models (GMMs) state distributions are used for classification. The appended jitter and shimmer features result in an increase in classification accuracy for several illustrative datasets, including the SUSAS dataset for human speaking styles as well as vocalizations labeled by arousal level for African elephant and Rhesus monkey species.

African elephant vocal communication I: antiphonal calling behaviour among affiliated females

African elephants, Loxodonta africana, are well known for their use of a low-frequency ‘rumble’ vocalization, which is thought to function in long-distance communication. Less work, however, has been conducted on short-distance communication within groups, and on spontaneously occurring vocal exchanges among identified individuals in particular. This is due in part to the fact that low-frequency rumbles are difficult to assign to individual callers. We collected vocal data on a group of six female African elephants housed at Disneys Animal Kingdom to determine whether they exchange rumbles in alternating sequences (also known as antiphonal calling). Subjects wore collars outfitted with microphones and radiotransmitters that allowed identification of individual callers, and behavioural and endocrine data were collected so that vocal activity could be examined in the context of social behaviour and reproductive state. First, we found that females did not produce rumbles at random, but were nearly twice as likely to produce rumbles shortly after rumbles from other group members. Second, the relative dominance rank and reproductive state of callers did not affect the probability of vocal response, but affiliative relationship with the caller had a strong influence on rumble response. Females were most likely to respond in kind to the rumbles of their most affiliated partners compared to less affiliated group members. Third, video analysis showed that rumble exchanges occurred in variable contexts, including when animals were out of contact, during reunions, and while in close proximity. Also, affiliated partners often vocalized in sequence when approached by dominant individuals. The results of these analyses show that affiliated female African elephants exchange rumbles antiphonally, and imply multiple functions for such vocal exchanges.

Bee Threat Elicits Alarm Call in African Elephants

Unlike the smaller and more vulnerable mammals, African elephants have relatively few predators that threaten their survival. The sound of disturbed African honeybees Apis meliffera scutellata causes African elephants Loxodonta africana to retreat and produce warning vocalizations that lead other elephants to join the flight. In our first experiment, audio playbacks of bee sounds induced elephants to retreat and elicited more head-shaking and dusting, reactive behaviors that may prevent bee stings, compared to white noise control playbacks. Most importantly, elephants produced distinctive “rumble” vocalizations in response to bee sounds. These rumbles exhibited an upward shift in the second formant location, which implies active vocal tract modulation, compared to rumbles made in response to white noise playbacks. In a second experiment, audio playbacks of these rumbles produced in response to bees elicited increased headshaking, and further and faster retreat behavior in other elephants, compared to control rumble playbacks with lower second formant frequencies. These responses to the bee rumble stimuli occurred in the absence of any bees or bee sounds. This suggests that these elephant rumbles may function as referential signals, in which a formant frequency shift alerts nearby elephants about an external threat, in this case, the threat of bees.

GPS determination of walking rates in captive African elephants (Loxodonta africana).

The movements of elephants in captivity have been an issue of concern for animal welfare activists and zoological professionals alike in recent years. In order to fully understand how movement rates reflect animal welfare, we must first determine the exact distances these animals move in the captive environment. We outfitted seven adult female African elephants (Loxodonta africana) at Disneys Animal Kingdom with collar-mounted global positioning recording systems to document their movement rates while housed in outdoor guest viewing habitats. Further, we conducted preliminary analyses to address potential factors impacting movement rates including body size, temperature, enclosure size, and social grouping complexity. We found that our elephants moved at an average rate of 0.409+/-0.007 km/hr during the 9-hr data collection periods. This rate translates to an average of 3.68 km traveled during the observation periods, at a rate comparable to that observed in the wild. Although movement rate did not have a significant relationship with an individuals body size in this herd, the movements of four females demonstrated a significant positive correlation with temperature. Further, females in our largest social group demonstrated a significant increase in movement rates when residing in larger enclosures. We also present preliminary evidence suggesting that increased social group complexity, including the presence of infants in the herd, may be associated with increased walking rates, whereas factors such as reproductive and social status may constrain movements.

African Elephant Alarm Calls Distinguish between Threats from Humans and Bees

The Samburu pastoralists of Northern Kenya co-exist with African elephants, Loxodonta africana, and compete over resources such as watering holes. Audio playback experiments demonstrate that African elephants produce alarm calls in response to the voices of Samburu tribesmen. When exposed to adult male Samburu voices, listening elephants exhibited vigilance behavior, flight behavior, and produced vocalizations (rumbles, roars and trumpets). Rumble vocalizations were most common and were characterized by increased and more variable fundamental frequencies, and an upward shift in the first [F1] and second [F2] formant locations, compared to control rumbles. When exposed to a sequence of these recorded rumbles, roars and trumpets, listening elephants also exhibited vigilance and flight behavior. The same behavior was observed, in lesser degrees, both when the roars and trumpets were removed, and when the second formants were artificially lowered to levels typical of control rumbles. The “Samburu alarm rumble” is acoustically distinct from the previously described “bee alarm rumble.” The bee alarm rumbles exhibited increased F2, while Samburu alarm rumbles exhibited increased F1 and F2, compared to controls. Moreover, the behavioral reactions to the two threats were different. Elephants exhibited vigilance and flight behavior in response to Samburu and bee stimuli and to both alarm calls, but headshaking behavior only occurred in response to bee sounds and bee alarm calls. In general, increasingly threatening stimuli elicited alarm calls with increases in F 0 and in formant locations, and increasing numbers of these acoustic cues in vocal stimuli elicited increased vigilance and flight behavior in listening elephants. These results show that African elephant alarm calls differentiate between two types of threat and reflect the level of urgency of threats.

Wild state secrets: ultra-sensitive measurement of micro-movement can reveal internal processes in animals

Assessment of animal internal “state” – which includes hormonal, disease, nutritional, and emotional states – is normally considered the province of laboratory work, since its determination in animals in the wild is considered more difficult. However, we show that accelerometers attached externally to animals as diverse as elephants, cockroaches, and humans display consistent signal differences in micro-movement that are indicative of internal state. Originally used to elucidate the behavior of wild animals, accelerometers also have great potential for highlighting animal actions, which are considered as responses stemming from the interplay between internal state and external environment. Advances in accelerometry may help wildlife managers understand how internal state is linked to behavior and movement, and thus clarify issues ranging from how animals cope with the presence of newly constructed roads to how diseased animals might change movement patterns and therefore modulate disease spread.

Male Masturbation in Free-Ranging Japanese Macaques

We studied masturbatory behavior of males over a 14-mo period in free-ranging Japanese macaques (Macaca fuscata yakui) on Yakushima island, Japan. We distinguished 2 types: (1) masturbation only and (2) masturbation with final ejaculation. Both types occurred in all 15 adult troop males. We calculated masturbation only as an index (M/h). It occurred over the whole year; however, it was more frequent during the mating than during the non-mating periods. Masturbation with ejaculation was restricted to the mating period. Male mating success correlated positively with social status. Both rate of masturbation only and masturbatory ejaculations increased as male social status and male mating success declined. Lower-ranking males had higher rates of masturbation only, i.e., they spent more time masturbating than higher-ranking males did. Lower-ranking males also had higher proportions of ejaculates for masturbation, while higher-ranking males used most of their ejaculates for mating.

Litter size and infant survivorship in wild groups of cotton-top tamarins (Saguinus oedipus) in Colombia.

Cotton‐top tamains (Saguinus oedipus) are a critically endangered primate found only in Colombia. Efforts to conserve this species are centered on developing effective management plans that integrate biological information regarding population dynamics and factors that influence their survival. This study documented infants born to wild cotton‐top tamarin females from 1994–2008 at two distinct field sites in northern Colombia. Our studies have shown that wild cotton‐top tamarins typically give birth to one litter each year and infant survival to 6 months of age was greater in the wild than has been reported in captive colonies. However, similar to reports from captive colonies, litter size of wild cotton‐top tamarins ranges from 1–3 infants, with twin litters most common. Here we report the first occurrence of triplet litters in nearly 20 years of observing wild cotton‐top tamarin groups. Over the first 3 months of life, wild‐born infants exhibited highest mortality during the first week of life, similar to reports from captive colonies. Infant survival in the wild also increases with successive litters as it does in captivity. However, inter‐birth interval, group size, and the number of adult males in the group did not appear to influence infant survival in the wild. The value of such long‐term data from field studies aids in the information that can be used to model future population trends and develop effective conservation plans for this critically endangered primate. Am. J. Primatol. 71:707–711, 2009.

INFANT AFRICAN ELEPHANT RUMBLE VOCALIZATIONS VARY ACCORDING TO SOCIAL INTERACTIONS WITH ADULT FEMALES

ABSTRACT Research on African elephant (Loxodonta africana) vocal communication has increased in recent years, yet there has been very little data collected on the vocal production of infant African elephants. Vocalizations were recorded from a group of five adult female African elephants and 3 dependent offspring (1 male and 2 female) at Disneys Animal Kingdom, Florida, U.S.A., using custom-designed audio-recording collars worn by the adult females. We measured both source and filter features of infant ‘rumble’ vocalizations made during affiliative social interactions and after cessation of nursing from adult females. Rumble vocalizations produced in the ‘nurse cessation’ context exhibited an upward shift in formant frequency locations, compared to rumbles produced during the ‘affiliation’ context. Additionally, call duration increased and fundamental frequencies decreased after nurse cessations for the male, but both females showed the opposite acoustic response. When infant rumbles accompanied nurse cessations, nursing was more likely to resume within 30 seconds compared to nurse cessations without vocalizations. These results suggest that infant rumbles associated with cessation of nursing reflect the motivational state of infants and may influence maternal responsiveness.