Heard but not seen: an acoustic survey of the African forest elephant population at Kakum Conservation Area, Ghana

This study, designed to survey forest elephants ( Loxodonta africana cyclotis ) at Kakum Conservation Area, Ghana, is the first to apply acoustic methods to elephant abundance estimation and to compare results with independent survey estimates. Nine acoustic sensors gathered sound continuously for 38 days. Low-frequency calling rates have been established as useful elephant abundance indices at a Namibian watering hole and a central African forest clearing. In this study, we estimated elephant population size by applying an abundance index model and detection function developed in central Africa to data from simultaneous sampling periods on Kakum sensors. The sensor array recorded an average of 1.81 calls per 20-min sampling period from an effective detection area averaging 10.27 km². The resulting estimate of 294 elephants (95% CI: 259–329) falls within confidence bounds of recent dung-based surveys. An extended acoustic model, estimating the frequency with which elephants are silent when present, yields an estimate of 350 elephants (95% CI: 315–384). Acoustic survey confidence intervals are at least half as wide as those from dung-based surveys. This study demonstrates that acoustic surveying is a valuable tool for estimating elephant abundance, as well as for detecting other vocal species and anthropogenic noises that may be associated with poaching.     

Visualizing Sound Emission of Elephant Vocalizations: Evidence for Two Rumble Production Types

Recent comparative data reveal that formant frequencies are cues to body size in animals, due to a close relationship between formant frequency spacing, vocal tract length and overall body size. Accordingly, intriguing morphological adaptations to elongate the vocal tract in order to lower formants occur in several species, with the size exaggeration hypothesis being proposed to justify most of these observations. While the elephant trunk is strongly implicated to account for the low formants of elephant rumbles, it is unknown whether elephants emit these vocalizations exclusively through the trunk, or whether the mouth is also involved in rumble production. In this study we used a sound visualization method (an acoustic camera) to record rumbles of five captive African elephants during spatial separation and subsequent bonding situations. Our results showed that the female elephants in our analysis produced two distinct types of rumble vocalizations based on vocal path differences: a nasally- and an orally-emitted rumble. Interestingly, nasal rumbles predominated during contact calling, whereas oral rumbles were mainly produced in bonding situations. In addition, nasal and oral rumbles varied considerably in their acoustic structure. In particular, the values of the first two formants reflected the estimated lengths of the vocal paths, corresponding to a vocal tract length of around 2 meters for nasal, and around 0.7 meters for oral rumbles. These results suggest that African elephants may be switching vocal paths to actively vary vocal tract length (with considerable variation in formants) according to context, and call for further research investigating the function of formant modulation in elephant vocalizations. Furthermore, by confirming the use of the elephant trunk in long distance rumble production, our findings provide an explanation for the extremely low formants in these calls, and may also indicate that formant lowering functions to increase call propagation distances in this species''.     

A TECHNIQUE TO ‘TAG’ SPECIFIC VOCALIZATIONS IN TAPE-RECORDINGS OF COMPLEX VOCAL INTERACTIONS

ABSTRACT

It has been suggested that African savanna elephants Loxodonta africana produce 31 different call types (Langbauer 2000). Various researchers have described these calls by associating them with specific behavioural contexts. More recently Leong et al. (2003) have attempted to classify elephant call types based on their physical properties. They classified 8 acoustically distinct call types from a population of captive elephants. This study focuses on one of these call types, the rumble, in a wild population of elephants in Kruger National Park, South Africa. A single family group of elephants was followed to record group behaviours and vocalizations from January through August 2001. By measuring the physical properties of 663 rumbles and subjecting these to cluster analysis, we present evidence that shows that rumbles can be categorized by their physical properties and that the resulting rumble types are associated with specific group behaviours. We characterize three types of rumbles that differ significantly by ten acoustic parameters. Two rumble types were associated with the elephant group feeding and resting, while the third was associated with socializing and agitation.     

The communicative content of the common marmoset phee call during antiphonal calling

Vocalizations are a dominant means of communication for numerous species, including nonhuman primates. These acoustic signals are encoded with a rich array of information available to signal receivers that can be used to guide species‐typical behaviors. In this study, we examined the communicative content of common marmoset phee calls, the species‐typical long distance contact call, during antiphonal calling. This call type has a relatively stereotyped acoustic structure, consisting of a series of long tonal pulses. Analyses revealed that calls could be reliably classified based on the individual identity and social group of the caller. Our analyses did not, however, correctly classify phee calls recorded under different social contexts, although differences were evident along individual acoustic parameters. Further tests of antiphonal calling interactions showed that spontaneously produced phee calls differ from antiphonal phee calls in their peak and end frequency, which may be functionally significant. Overall, this study shows that the marmoset phee call has a rich communicative content encoded in its acoustic structure available to conspecifics during antiphonal calling exchanges. Am. J. Primatol. 72:974–980, 2010. © 2010 Wiley‐Liss, Inc.     

Complex Vocal Responses to Conspecific Calls in Whistling Frogs Eleutherodactylus johnstonei: Playback Experiments in the Field

We simulated the presence of an acoustic competitor by broadcasting conspecific playbacks to males of Johnstone's whistling frog, Eleutherodactylus johnstonei, in the field. We broadcast calls that differed in duration (short, typical, and long), dominant frequency (high, typical, and low), and period (short, typical, and long), and analyzed male vocal responses. We tested the hypothesis that males respond by escalating vocally when they are exposed to female‐attractive calls and by ignoring unattractive ones. At the population level, males responded to playbacks in ways that would potentially increase their attractiveness with regard to solo calling: males increased the duration, reduced the dominant frequency, and increased their calling effort (duty cycle), despite an increase in call period. The modification of call duration occurred only in response to playbacks of low‐frequency calls, long calls, and short‐period calls (selective response), while the modification of the dominant frequency was independent of the characteristic of the playback (fixed response). Contrary to the expected, males did not reduce the call period when they were exposed to attractive playbacks. At the ultimate level, the results suggest energy‐saving strategies. In addition, males seem to trade off call period for the avoidance of acoustic interference with attractive calls as calling effort was typically increased by increasing call duration but only rarely by reducing the call period. Interactive playbacks are necessary to better understand the calling strategies of males of E. johnstonei.     

Call-type matching in vocal exchanges of free-ranging resident killer whales, Orcinus orca

Previous sound recordings of resident (fish-eating) killer whale groups have revealed matrilineal group-specific call repertoires and a strong tendency for calls of the same type to be produced in series. Vocal interactions between individual free-ranging animals, however, have remained unexplored because it has not been possible to identify signallers reliably with a single hydrophone. Here we link acoustic arrivals of calls on a towed hydrophone array with visual tracking of photo-identified individuals to ascribe calls to a focal animal when it was separated from other members of its matrilineal group by more than 35 m, and thereby out of visual range. We confirm that individual members of a matrilineal group share a repertoire of stereotyped calls, and we statistically examine timing of stereotyped calls produced by one individual relative to calls produced by other members of its group. Analysis of the intervals between stereotyped calls indicated that calls were produced in group bouts with a criterion interval of 19.6 s separating bouts. We were therefore careful to develop randomization tests that preserved call interval structure. Focal whales produced 36% of their calls within 5 s of a call from a nonfocal animal, four times more calls than expected by chance based upon a rotation randomization test. Consecutive calls produced by different individuals during group-calling bouts matched call type more than expected by chance. Vocal exchanges of stereotyped calls with type matching appear to be an important aspect of intragroup calling in killer whales, although the function of this calling behaviour remains to be explored.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

Long Recording Sequences: How to Track the Intra-Individual Variability of Acoustic Signals

Recently developed acoustic technologies - like automatic recording units - allow the recording of long sequences in natural environments. These devices are used for biodiversity survey but they could also help researchers to estimate global signal variability at various (individual, population, species) scales. While sexually-selected signals are expected to show a low intra-individual variability at relatively short time scale, this variability has never been estimated so far. Yet, measuring signal variability in controlled conditions should prove useful to understand sexual selection processes and should help design acoustic sampling schedules and to analyse long call recordings. We here use the overall call production of 36 male treefrogs (Hyla arborea) during one night to evaluate within-individual variability in call dominant frequency and to test the efficiency of different sampling methods at capturing such variability. Our results confirm that using low number of calls underestimates call dominant frequency variation of about 35% in the tree frog and suggest that the assessment of this variability is better by using 2 or 3 short and well-distributed records than by using samples made of consecutive calls. Hence, 3 well-distributed 2-minutes records (beginning, middle and end of the calling period) are sufficient to capture on average all the nightly variability, whereas a sample of 10 000 consecutive calls captures only 86% of it. From a biological point of view, the call dominant frequency variability observed in H. arborea (116Hz on average but up to 470 Hz of variability during the course of the night for one male) challenge about its reliability in mate quality assessment. Automatic acoustic recording units will provide long call sequences in the near future and it will be then possible to confirm such results on large samples recorded in more complex field conditions.     

SOCIETY FOR EXPERIMENTAL BIOLOGY CONFERENCE,UNIVERSITY OF ST. ANDREWS,APRIL 1995

We describe the interactive nature of vocalizations emitted by African elephant (Loxodonta africana) family groups while visiting waterholes. Despite being in close visual contact with each other, individuals called interactively within a series of overlapping or antiphonal vocal bouts that increased significantly while departing from the waterhole. After initiating departure from the waterhole, elephants in this study increased their calling rate and their production of overlapping vocal bouts. The majority of calls either overlapped or occurred within 1.5 seconds of another call as part of an antiphonal bout, rather than as isolated calls. The departure of larger herds was accompanied by a greater number of calling bouts. The increase in interactive antiphonal bouts during departure might serve to facilitate group coordination and cohesion, as well as possibly reinforce social bonds. The longer repeated bouts could also facilitate eavesdropping by distant elephants by boosting signal detection since the repetition of these longer calls may yield an increase in the signal-to-noise ratio (SNR) that reduces the noise floor for better longer distance communication.     

Extending bioacoustic monitoring of birds aloft through flight call localization with a three‐dimensional microphone array

Bioacoustic localization of bird vocalizations provides unattended observations of the location of calling individuals in many field applications. While this technique has been successful in monitoring terrestrial distributions of calling birds, no published study has applied these methods to migrating birds in flight. The value of nocturnal flight call recordings can increase with the addition of three‐dimensional position retrievals, which can be achieved with adjustments to existing localization techniques. Using the time difference of arrival method, we have developed a proof‐of‐concept acoustic microphone array that allows the three‐dimensional positioning of calls within the airspace. Our array consists of six microphones, mounted in pairs at the top and bottom of three 10‐m poles, arranged in an equilateral triangle with sides of 20 m. The microphone array was designed using readily available components and costs less than $2,000 USD to build and deploy. We validate this technique using a kite‐lofted GPS and speaker package, and obtain 60.1% of vertical retrievals within the accuracy of the GPS measurements (±5 m) and 80.4% of vertical retrievals within ±10 m. The mean Euclidian distance between the acoustic retrievals of flight calls and the GPS truth was 9.6 m. Identification and localization of nocturnal flight calls have the potential to provide species‐specific spatial characterizations of bird migration within the airspace. Even with the inexpensive equipment used in this trial, low‐altitude applications such as surveillance around wind farms or oil platforms can benefit from the three‐dimensional retrievals provided by this technique.     

Neotropical frogs and mating songs: The evolution of advertisement calls in glassfrogs

Anurans emit advertisement calls with the purpose of attracting mates and repelling conspecific competitors. The evolution of call traits is expected to be associated with the evolution of anatomical and behavioural traits due to the physics of call emission and transmission. The evolution of vocalizations might imply trade‐offs with other energetically costly behaviours, such as parental care. Here, we investigated the association between body size, calling site, parental care and call properties (call duration, number of notes, peak frequency, frequency bandwidth and call structure) of the advertisement calls of glassfrogs (Centrolenidae)—a family of Neotropical, leaf‐dwelling anurans—using phylogenetic comparative methods. We also explored the tempo and mode of evolution of these traits and compared them with those of three morphological traits associated with body size, locomotion and feeding. We generated and compiled acoustic data for 72 glassfrog species (46% of total species richness), including representatives of all genera. We found that almost all acoustic traits have significant, but generally modest, phylogenetic signal. Peak frequency of calls is significantly associated with body size, whereas call structure is significantly associated with calling site and paternal care. Thus, the evolution of body size, calling site and paternal care could constrain call evolution. The estimated disparity of acoustic traits was larger than that of morphological traits and the peak in disparity of acoustic traits generally occurred later in the evolution of glassfrogs, indicating a historically recent outset of the acoustic divergence in this clade.     

The Vocal Repertoire of Barbary Macaques: A Quantitative Analysis of a Graded Signal System

The vocal behavior of Barbary macaques (Macaca sylvanus) was studied in an outdoor enclosure at Rocamadour, France. Ad libitum recordings were made across a broad array of socioecological contexts from 92 individually identified subjects from all age—sex classes. From the recordings, 8479 calls were sampled and submitted to a Fourier transform. A custom software program was used to determine 35 acoustic parameters describing the call in terms of its frequency and time dynamics. On these parameters a cluster analysis was used to examine the acoustic morphology of the Barbary macaque vocal repertoire. The analysis revealed a highly graded structure with intergraded variations between different clusters (‘call types’). There were clear age-related preferences in usage of different clusters, but animals of all ages were potentially able to produce the whole array of call types. The major sex difference was a differential use of certain call types in specific situations. Despite a clear sex dimorphism in body size no significant sex differences in the acoustic structure of calls in adults was found. There were no unidirectional relationships between the acoustic structure of a call and the context in which it was uttered, although call usage was more specific in some contexts than in others. The results suggest that the major factor underlying the variation in acoustic structure reflect the internal state of the caller.     

The units of perception in the antiphonal calling behavior of cotton-top tamarins (Saguinus oedipus): playback experiments with long calls

We investigated how the acoustic structure of the cotton-top tamarin monkey's (Saguinus oedipus) combination long call relates to the antiphonal calling behavior of conspecifics. Combination long calls can function as contact calls and are produced by socially isolated individuals. Often conspecifics respond to these calls with their own long calls. Structurally, these calls are always composed of one or more 'chirps' followed by two or more 'whistles'. We compared the antiphonal calling responses to playbacks of complete, naturally produced long calls versus single whistles or single chirps. Subjects responded significantly more to whole calls than to either syllable-type alone. Thus, our data suggest that, in terms of the antiphonal calling behavior of socially isolated conspecifics, the whole long call is the unit of perception.     

沼水蛙繁殖期鸣声特征及鸣叫节律

鸣叫对无尾两栖类动物的生存与繁殖起重要作用。蛙类的鸣叫行为受到环境因素影响表现出一定的节律性。2016年8和9月,采用录音机和指向性话筒,在野外录制了57只沼水蛙（Hylarana guentheri）的鸣声并对其鸣声特征进行分析;通过悬挂录音笔和自动温湿度记录仪研究了沼水蛙鸣叫节律（17 d）及其与环境温度、相对湿度的关系。结果显示,沼水蛙的鸣声由1 ~ 4个音节组成,不同类型鸣声间的音节主频、音节时长存在显著差异（P < 0.05）。该物种全天具有鸣叫行为,13:00 ~ 14:00时为鸣叫高峰期。白天单音节鸣声、双音节鸣声、三音节鸣声、总鸣声和总音节的数量较夜晚显著增加（P < 0.01）。鸣声数量和音节数量均与环境温度呈正相关（P < 0.01）。结果表明,沼水蛙通过改变音节数量、音节主频和音节时长改变鸣叫策略。沼水蛙的鸣叫行为具有昼夜节律性且受环境温度的影响。     

Female call preferences in tree-hole frogs: why are there so many unattractive males?

In tree-hole frogs, Metaphrynella sundana, the fundamental call frequency varies widely between males. In field playback experiments, females strongly preferred calls from the lower range of frequencies found in the population. There was no correlation, however, between male size and call frequency, as is normally the case for anurans, so large males were not necessarily more attractive to females. Presence or absence of upper harmonics in the call had no effect on female choice. Tree holes with shallow air columns were more often used by calling frogs, and were presumably more common, than deep holes. Since male M. sundana actively exploit the resonant properties of tree holes for mate attraction, and high frequencies match comparatively shallow holes, the benefits of attaining acoustic matching probably select for high-frequency calls. In addition, males with high-frequency calls may be heard from a greater distance in the vicinity of torrent streams. Since the level of such noise in the forest varies in time and space, different frequencies may prove optimal in different contexts, thereby preserving the observed variation within the population. Having an ‘unattractive’ high-frequency call should be potentially beneficial only when calling males do not congregate, a condition that our data suggest is fulfilled in this system.     

Acoustic estimation of wildlife abundance: methodology for vocal mammals in forested habitats

Habitat loss and hunting pressure threaten mammal populations worldwide, generating critical time constraints on trend assessment. This study introduces a new survey method that samples continuously and non‐invasively over long time periods, obtaining estimates of abundance from vocalization rates. We present feasibility assessment methods for acoustic surveys and develop equations for estimating population size. As an illustration, we demonstrate the feasibility of acoustic surveys for African forest elephants (Loxodonta africana cyclotis). Visual surveys and vocalizations from a forest clearing in the Central African Republic were used to establish that low‐frequency elephant calling rate is a useful index of elephant numbers (linear regression P < 0.001, r_adj.² = 0.58). The effective sampling area was 3.22 km² per acoustic sensor, a dramatic increase in coverage over dung survey transects. These results support the use of acoustic surveys for estimating elephant abundance over large remote areas and in diverse habitats, using a distributed network of acoustic sensors. The abundance estimation methods presented can be applied in surveys of any species for which an acoustic abundance index and detection function have been established. This acoustic survey technique provides an opportunity to improve management and conservation of many acoustically‐active taxa whose populations are currently under‐monitored.     

Repeated call sequences and behavioural context in long-finned pilot whales off Cape Breton,Nova Scotia,Canada

Repeated calls are part of the vocal repertoire of a diverse array of species, often presented in sequences that take time and effort on the part of the signal producer. Rhythmic repeated call sequences make up a significant portion of long-finned pilot whale (Globicephala melas) vocal production, yet the function of these sequences has not been investigated until now. In this study, we explored the relationship between behavioural context and the presence of these vocal sequences using recordings of a population of pilot whales found off Cape Breton, Nova Scotia, Canada. We used a binomial logit-link generalized linear model to look for possible predictors of the presence of repeated call sequences. They were more common in recordings of socializing whales than in those of whales in other behavioural states, and least common in resting whales. These vocal repetitions were also more common with larger group size. These results suggest that sequences function in maintaining contact and cohesion within this social species, possibly also serving in individual or group identification. The context of repeated call sequences indicate that they are not primarily mother–calf interactions, as they are heard just as commonly from groups without young. Future studies of pilot whale repeated call sequences should include individual-level behaviour and detailed acoustic calling context.     

Are acoustical parameters of begging call elements of thin-billed prions related to chick condition?

Chicks of burrowing petrels use begging calls to advertise their hunger levels when parents arrived at the nest. In a previous study, adult thin-billed prions Pachyptila belcheri responded to higher begging call rates of their single chick by regurgitating larger meals. We tested whether acoustic parameters of begging call elements may also be involved in signalling. To describe variation in begging, we determined begging session parameters, namely the duration, number of calls and the mean and maximum rate of calling. We then digitised calls and carried out a semi-automatic extraction of six acoustic parameters of call elements, including mean and maximum acoustic frequency, the length of call elements and the location of the maximum frequency and amplitude within calls. Chicks showed strong individual differences in all parameters. While the session parameters were correlated with body condition and with the meal size the chick received, none of the acoustic parameters were related to body condition and provisioning. A cross-fostering experiment showed the same pattern, as only session parameters changed related to an experimentally altered body condition, while acoustical cues appear to play no role in signalling hunger levels. We suggest that this may be explained by the absence of sibling competition in these birds. As parents do not need to decide which chick to feed, immediate information on condition at the time of adult arrival may not be required.     

Social context and prey composition are associated with calling behaviour in a diving seabird

Social cohesion and prey location in seabirds are largely enabled through visual and olfactory signals, but these behavioural aspects could potentially also be enhanced through acoustic transfer of information. Should this be the case, calling behaviour could be influenced by different social–ecological stimuli. African Penguins Spheniscus demersus were equipped with animal-borne video recorders to determine whether the frequency and types of calls emitted at sea were dependent on behavioural modes (commuting, sedentary and dive bout) and social status (solitary vs. group). For foraging dive bouts we assessed whether the timing and frequency of calls were significantly different in the presence of schooling prey vs. single fish. The probability of call events was significantly more likely for birds commuting early and late in the day (for solitary birds) and during dive bouts (for groups). During foraging dive bouts the frequency of calls was significantly greater for birds diving in the presence of schooling fish and birds called sooner after a catch in these foraging scenarios compared with when only single fish were encountered. Three call types were recorded, 'flat', 'modulated' and 'two-voice' calls, but there was no significant relationship detected with these call types and behavioural modes for solitary birds and birds in groups. The results of this study show that acoustic signalling by African Penguins at sea is used in a variety of behavioural contexts and that increased calling activity in the presence of more profitable prey could be of crucial importance to seabirds that benefit from group foraging.