Int J Primatol (2008) 29:615625

DOI 10.1007/s10764-008-9253-x

Orangutan Long Call Degradation

and Individuality Over Distance:
A Playback Approach
Adriano R. Lameira & Serge A. Wich

Received: 6 May 2007 / Accepted: 5 February 2008 /

Published online: 3 June 2008
# Springer Science + Business Media, LLC 2008

Abstract Researchers have documented individual vocal recognition in several

primate species but do not know whether the changes in acoustical parameters that
might occur over distance influence the informational content of a call that relates to
individuality. Accordingly, we performed playback experiments using male orangutan
long-distance calls (long calls) and rerecorded them at increasing distances from the
source. We aimed to determine 1) which acoustical parameters changed over distance
and 2) whether the percentage of calls that a discriminant analyses would assign to the
correct individual would change over distance. High-frequency harmonics were
attenuated and lost with increasing distance, but other parameters did not change. The
percentage of calls assigned to the correct individual did not change over distance,
indicating that even though there are some acoustical changes over distance the
opportunity for other individuals to recognize the caller remains similar until 300 m,
which was the maximum distance at which we rerecorded calls. Extending similar
experiments to other primate species and other taxa, while subsequently conducting
experiments to assess whether individual discrimination by receivers is indeed based
on relatively stable acoustical parameters, would forward our understanding of
acoustic communication.
Keywords Pongo pygmaeus wurmbii . sound distortion . vocal discrimination .
vocalization

A. R. Lameira
Behavioral Biology Group, University of Utrecht, 3508TB Utrecht, The Netherlands
S. A. Wich (*)
Great Ape Trust of Iowa, Des Moines, IA 50320, USA
e-mail: swich@greatapetrust.org
S. A. Wich
Behavioral Biology Group, University of Utrecht, 3508 TB Utrecht, The Netherlands

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A.R. Lameira, S.A. Wich

Introduction
All primate species live in social organizations in which individual recognition is
important for aspects of daily life such as mate choice, alliance and coalition
formation, foraging, and dominance hierarchies. Though visual recognition is
important at short distances (Dominy et al. 2001), there is sufficient evidence that
individual recognition on the basis of acoustical characteristics occurs in many
primate species for both short- and long-distance vocalizations (Cheney and Seyfarth
1980, 1999; Fischer 2004; Rendall et al. 2004).
Bioacoustical researchers show that vocalizations may change from its source to
its receiver as a function of attenuation, random amplitude fluctuations, and
reverberations, i.e., distortion (Brown 2003; Marten et al. 1977; Waser and Brown
1986; Waser and Waser 1977; Wiley and Richards 1978). The vocalizations of many
species are therefore thought to have been selected to reduce the impact of the
aforementioned factors. However, with increasing distance, distortion is inevitable
and it is informative to examine how such distortion influences the potential for
individual recognition of vocalizations by receivers.
The importance of distortion is especially apparent for long-distance vocalizations (Waser and Waser 1977). An important, but as far as we know neglected
question in primate studies, is whether distortion poses a constraint on individual
discrimination as distance between the signaler and the receiver increases (Brown
2003). Individual recognition of long calls is essential to explain the results of
studies that indicate that males and females respond differentially to long calls of
different males (Delgado 2003; Mitani 1985; Mitra Setia and van Schaik 2007).
Such studies are based on the assumption that the receiver can individually
distinguish long calls at a certain distance. However, it remained unknown whether
distortion poses a constraint on individual discrimination. We aimed to answer the
question experimentally.
Long calls are the most prominent and most studied orangutan vocalization
(Davila Ross 2003; Davila Ross and Geissmann 2007; Delgado 2003, 2007;
Galdikas 1983; Hardus et al. in press; Mitani 1985). Receivers can hear long calls up
to several hundred meters. They consist mainly of repeated sequences that might
differ in their acoustical structure (Davila Ross and Geissmann 2007; Rijksen 1978).
Researchers believe that long calls mediate dominance relationships among adult
males (Delgado and van Schaik 2000; Galdikas 1983; MacKinnon 1974; Singleton
and van Schaik 2002), attract adult females from a long distance to facilitate mating
(Galdikas 1983; MacKinnon 1974, 1979; Mitani 1985; Schrmann and van Hooff
1986), and act as a mechanism via which individuals maintain loose associations
within a dispersed community and as a coordinating signal for seasonal movements
of whole communities (cf. Delgado and van Schaik 2000). For all of the functions,
individual recognition of long calls is important. The fact that long calls show
acoustically individual differences is an indication that individual recognition is
likely (Delgado 2007).
Therefore, we aimed to examine experimentally 1) whether acoustic parameters
in orangutan long calls change with increasing distance from the playback speaker
and 2) whether individual classifications of long calls become less accurate with
increasing distance from the playback speaker. A long call consists of a bubble-

Orangutan Long Call Degradation

617

like introduction that leads to a relatively long sequence of bark-like sounds:

pulses (Fig. 1), and ends with a series of sigh-like sounds (Galdikas 1983, Mitani
1985). The acoustic parameters we used for the purpose are: 1) pulse duration, 2) F0
begin frequency, 3) F0 high frequency, 4) F0 max frequency, 5) frequency
bandwidth, and 6) Fn max frequency (Fig. 1). We based our choice of parameters
on recent studies on orangutans showing them to be important in individual
discrimination of long calls (Davila Ross and Geissmann 2007; Delgado 2007).
Therefore, we were most interested in the stability of the parameters over distance.
Pulse duration (s) is the time interval between the beginning and the end of one
pulse. F0 begin frequency (Hz) is the frequency with the highest energy at the
beginning of the pulses F0, i.e., the highest amplitude frequency in rectangle 2 in
Fig. 1. F0 (fundamental frequency) is the first and lowest harmonic of one pulse and
typically the one with the highest amplitude. F0 high frequency (Hz) is the frequency
with the highest energy at the instant F0 reaches its highest point, i.e., the highest
amplitude frequency in rectangle 3 in Fig. 1. F0 max frequency (Hz) is the frequency
with the highest energy emitted in a pulses F0, independently from its location in
the pulse, i.e., the highest amplitude frequency in rectangle 4 in Fig. 1. Frequency
bandwidth (Hz) is the frequency interval between the lowest frequency at F0
and the highest frequency in the pulse at Fn. Fn was the highest harmonic
recognizable and was not necessarily the same over the pulses of one call. Fn max
frequency (Hz) is the frequency with the highest energy in the highest harmonic
recognizable.
Based on the literature (Brown 2003; Hunter and Krebs 1979; Konishi 1970;
Morton 1975), we predicted that pulse duration should not change over distance
because it is based on a low-frequency part of the pulse, which is not likely to
reverberate. Because the obstacles in a forest are relatively small in relation to the
wavelength of low-frequency signals, we expected that F0 begin frequency, F0 high
frequency, and F0 max frequency might change only slightly with distance, but that
the high-frequency parameters of the call (frequency bandwidth and Fn max

Fig. 1 Spectrogram showing 1) pulse duration, 2) F0 begin frequency (measurement window 70 ms), 3)
F0 high frequency (measurement window 70 ms), 4) F0 max frequency (measurement window as long as
duration of F0), 5) frequency bandwidth, and 6) Fn max frequency (measurement window as long as
duration of Fn). A total of 3 pulses are shown.

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A.R. Lameira, S.A. Wich

frequency) will be more distorted over distance (Brown 2003). Because individual
recognition of long calls over distance is likely to be very important in orangutan
social organization we predicted that individual classification would remain similar
with increasing distance.

Methods
Study Site
We conducted the study from December 2004 to September 2005 at Tuanan, Central
Kalimantan, Indonesia. The site (209S; 11426E) consists of forests on shallow
peat, of varying thickness, ca. 2 m. Research on the wild orangutans in Tuanan
started in January 2003, and we recognized 36 individuals, 9 of which were flanged
males.
Data Collection, Playback Studies, and Data Analysis
We opportunistically recorded long calls at 1015 m from vocalizing males via a
Marantz Analogue Recorder PMD222 in combination with a Sennheiser Microphone ME 64 or a Sony Digital Recorder TCD-D100 in combination with a Sony
Microphone ECM-M907. We included no long calls produced toward observers. We
recorded a total of 49 long calls (699 pulses from 7 males, with a range of 242
pulses per long call) from which we selected the best quality recordings. The lower
quality recordings were not from one particular context, so we believe our
recordings are representative of all long calls that the orangutan males produced.
We subsequently digitized them at 44.1 kHz, amplified them to 30.50.5 kU (the
amplitude unit used by Raven Software), and filtered them >2.250.1 kHz via
Raven Interactive Sound Analysis Software (2003, Cornell Lab of Ornithology,
Ithaca, NY). We excised background noise that was very clear, e.g., bird songs, but
in reality it was necessary only in 2 of the playback long calls and did not involve
the actual pulses and therefore did not influence the pulse characteristics. We
conducted amplification and filtering to standardize long call amplitude for
playbacks. For each male, we selected several representative pulses from the
beginning and middle of high-quality long calls and then transferred them from the
computer to a TCD-D100 digital Sony recorder. Pulses at the end of long calls
were often less clear and therefore discarded. In addition, Delgado (2003)
suggested that long calls convey information in the early part of the long call. We
played back the pulses in sequences of pulses for each male to mimic natural pulse
series in long calls from a forest platform 10 m above the ground, via the TCD-D100
in combination with a Nagra DSM monitor that always faced north. Peak sound
pressure levels (SPL) for all playbacks was 1005 dB at 1 m from the loudspeaker,
which equals the natural SPL of orangutan long calls at Tuanan (Lameira and Wich,
unpubl. data). We rerecorded the played back long calls via a Marantz PMD222
analog recorder with a Sennheiser ME 64 microphone at a height of 1.50.1 m. We
acknowledge that conducting the playback and recording the playback at only 1
height at 1 location is a limitation to generalizing the results of our experiments. But

Orangutan Long Call Degradation

619

conducting a large set of playbacks from different heights and making recordings
at different heights would have greatly disturbed the orangutans in the area.
Therefore we restricted ourselves to a limited set of experiments. However, the
forest in the area appears very homogenous. We made recordings of the same
played back pulses at 6 different distances (50 m, 100 m, 150 m, 200 m, 250 m,
and 300 m) in a straight line from the speaker and always at the same place for
each call. At >300 m the long call was too faint to record. We selected the transect
along which we recorded the experimental long calls based on characteristics that
are representative of the forest in the area. Therefore the forest along the transect
included 2 minor gaps in the canopy because they are a common feature of peat
swamp forests in the area.
The terrain where we conducted the experiments was flat and we conducted
experiments between 0530 h and 0630 h to minimize amplitude fluctuations due to
atmospheric turbulence (Richards and Wiley 1980). There was no wind during any
of the experiments, and we conducted all experiments during dry mornings when
there had been no rainfall since the previous night.
We used 11 long calls from 7 of the 9 identified males, in a total of 110 pulses
(23, 6, 20, 20, 15, 5, 21) for the playback experiments. We transformed long call
recordings into spectrograms (window type = Hanning; spectrogram configuration:
time grid spacing = 256; samples/frame overlap = 50%; frequency grid spacing =
86.1; window size = 512 samples; 3-dB bandwidth = 124 Hz, i.e., narrow-band like
spectrogram; Fig. 1). We measured the values of each parameter in the spectrograms
of each distance rerecording.
We conducted the discriminant analyses via SPSS 11.0 to examine differences
between individuals and followed similar methods of other studies that have
examined individual variation in vocalizations (Wich et al. 2003). The probability of
correctly assigning a call to the correct individual by chance was 0.14 (1/7 males).
We validated call assignment via discriminant analyses with a leave-one-out
procedure, which followed methods similar to those of Wich et al. (2003). We used
2-tailed Spearman correlations to confirm if any parameter and/or percentage of
correct assignments correlates with distance. As the unit of analyses we used the
averages of the average measurement values on the pulses for each male to avoid
overrepresenting certain males. We corrected p values via a sequential Bonferroni
procedure, except where mentioned otherwise (Hochberg 1988).

Results
Long Call Distortion Over Distance
The mean values of each parameter varied greatly over distance, but few declined
significantly with increased distance (Fig. 2). Pulse duration, F0 max frequency, F0
begin frequency, and F0 high frequency did not significantly change with increasing
distance (Spearman correlation coefficient: r=0.14, p=0.79; r=0.83, p=0.17;
r=0.49, p=0.99; r=0.43, p=0.79, respectively, n=6 for all tests). The only 2
parameters that showed a significant decline with distance are frequency bandwidth

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A.R. Lameira, S.A. Wich

Fig. 2 Parameters mean per distance and regression lines. (a) Pulse duration. (b) F0 begin frequency. (c)
F0 high frequency. (d) F0 max frequency. (e) Fn max frequency. (f) Frequency bandwidth.

and Fn max frequency (Spearman correlation coefficient: r=1.0, p<0.01; r=1.0,

p<0.01, respectively, n=6 for both tests).
Long Call Discrimination Over Distance
We first conducted a discriminant analysis and leave-one-out validation employing
all parameters, which for the original recordings yielded a correct classification of

Orangutan Long Call Degradation

621

Table 1 Percentages of variance explained per canonical discriminant function used in discriminant
analysis employing all the parameters, for the 6 distances
Distance (m)/Function

1
2
3
4
5
6
a

0 ma

50 m

100 m

150 m

200 m

250 m

300 m

61.2
23.7
12.1
2.8
0.2
0.0

55.4
22.0
17.3
2.5
2.2
0.5

59.4
25.7
13.3
1.4
0.2
0.0

58.8
18.5
14.1
5.1
3.4
0.0

76.4
14.2
4.2
4.1
1.1
0.0

72.5
12.6
8.8
4.7
1.2
0.2

67.5
17.1
9.4
5.4
0.6
0.0

0 meter indicates the original recordings.

41.4% of the long calls. Via the cross validation procedure, the correct classification
is reduced to 39.9%. The first and second discriminant functions explained 84.9% of
the variance (Table 1). At 50 m from the speaker, the first 2 functions explained
77.4% (Table 1). The percentage of calls assigned to the correct individual at this
distance is 45.5% (31.8% with leave-one-out validation). We conducted a similar
analysis for each distance and found that the correct assignments varied between
35.4% and 50.0% (20.2% and 32.8% with the leave one out validation, Fig. 3). The
percentage of correct assignments showed no correlation with distance (Spearman
correlation coefficient, 2-tailed: r=0.09, p=0.82; with leave-one-out validation, 2tailed: r=0.43, p=0.40, Table 2).
We repeated the discriminant, leave-one-out, and Spearman correlation analyses
for each parameter separately to evaluate the importance in the discrimination of
each parameter (Table 2). The parameter with the highest percentage for correctly
assigning calls is pulse duration. All the frequency parameters showed lower
percentages (Table 2). The percentage of correct assignments does not correlate
significantly with distance for any of the parameters (Table 2; Fig. 4). The leaveone-out validation procedure yielded similar results (Table 2; Fig. 4). Inspecting
Fig. 4 also indicates that pulse duration is the parameter that over distance has the

Fig. 3 (a) Percentage of calls assigned correctly versus distance for all 6 parameters combined. (b) The
same analysis but with leave-one-out validation. The horizontal dashed line represents the 14% chance
level.

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A.R. Lameira, S.A. Wich

Table 2 Results of the discriminant analyses with and without leave-one-out validation, for all parameters
combined and for each parameter separately
Parameters

min-max%a min-max%b SEa

SEb

6 parameters
combined
Pulse duration
F0 begin freq.
F0 high freq.
F0 max freq.
Bandwidth
Fn max freq.

35.450.0

20.232.4

2.28 5.99 0.09 0.88 0.26 0.62 41.4

39.9

22.925.5
18.222.7
17.324.8
15.725.5
14.328.2
12.129.1

21.824.5
10.022.7
16.423.8
6.419.8
10.928.2
11.029.1

0.46
0.74
1.16
1.57
2.09
2.29

40.3
27.7
28.3
23.6
23.6
27.5

0.48
5.28
3.49
5.12
2.66
2.77

ra

0.44
0.60
0.09
0.26
0.71
0.83

pa

1.0
0.84
0.87
1.0
0.55
0.24

rb

0.84
0.49
0.43
0.31
0.6
0.89

pb

0.20
1.0
0.80
0.54
0.84
0.12

Total
Total
samplea sampleb

41.6
27.7
28.3
23.6
23.6
27.5

Discriminant analysis.
Leave-one-out validation.
min-max% = minimal and maximal percentage of correctly assigned calls; SE = standard error; r = r value
from Spearmans correlation coefficient, 2-tailed; p = p value of Spearman test after a Bonferroni
correction.

most stable percentage of correctly assigned calls, whereas the other parameters
either fluctuated widely or decreased (Fig. 4).

Discussion
We aimed to address whether acoustical parameters of orangutan long calls changed
with distance from the source and whether this affected potential individual
discrimination. Pulse duration is the parameter that overall yielded the highest
percentages of correctly assigned calls (Table 2). Ambient factors, e.g., branches,
leaves, trunks, or atmospheric factors, produce rapid attenuation of high frequencies
and a shift to lower frequencies (Brown 2003; Konishi 1970; Wiley 1991).

Fig. 4 (a) Percentage of correctly assigned calls versus distance for all parameters separately and
regression lines. (b) Same analyses, but with leave-one-out validation. The horizontal dashed line
represents the 14% chance level.

Orangutan Long Call Degradation

623

Accordingly, bandwidth and Fn max frequency decreased significantly with distance.

Therefore, our results support the prediction that high-frequency parameters are
more distorted with distance than low-frequency parameters are.
Thus pulse duration and the 3 low-frequency parameters are expected to be the
best call parameters for information coding over distance because they show the
least change with distance, which is consistent with the findings of Delgado (2003).
Delgados data (2003) also suggest that frequency bandwidth and Fn max frequency
are parameters that discriminate long calls correctly. However, Delgado used only
calls that he recorded rather close to the caller. In fact, as we show (Fig. 4), at close
distances frequency bandwidth and Fn max frequency perform well as discriminant
parameters, but thereafter drop considerably.
Our results indicate that the percentage of long calls correctly assigned to an
individual remained relatively similar with increasing distance. The percentage of
correct assignments was somewhat higher at 150 m and 250 m, which could be
related to 2 gaps occurring at there: there may have been less influence of vegetation
on sound transmission and therefore less distortion of the long calls, enabling a more
accurate measure of the acoustic parameters and consequently higher percentages of
correct assignments via discriminant analyses. It is important to note that gaps are a
common feature of peat swamp forests in the area and thus do not decrease the
ecological validity of our experiment. It would be very interesting to conduct a
similar set of experiments in dryland forests where canopy structure differs. Because
there is no comparative forest canopy structure study between the peat swamp and
dryland forests, we cannot make specific predictions as to how long call distortion
might differ between the 2 forest types.
The overall percentages of correct assignment for the playback long calls show
considerable overlap with values in other studies, e.g., 2188% per parameter for
calls from several individuals (Delgado 2007). The results indicate that at 300 m
receivers can potentially still distinguish individuals via their long calls at a
discriminant level similar to that of the original calls. Though we attempted to record
the playbacks >300 m from the source, the decrease in amplitude over distance did
not allow for a high enough amplitude to make proper recordings at 350 m. Thus,
though frequency parameters degrade over distance, individuality of the long calls
remains similar with increasing distance until 300 m. In the wild orangutan, long
calls are audible at >300 m, but Mitra Setia and van Schaik (2007) showed that
females react only to callers <400 m away. Moreover, they showed that females
differentiate their responses depending on which male emits the long call. Their
results indicate that females can differentiate between male long calls only when
they are emitted <400 m away, which corresponds with our results that indicate at
300 m individual assignments of long calls remain high. The fact that we could
make no recording at >300 m and that long calls are audible from longer distances
might indicate not only that the power of the speaker is not similar to that of an
orangutan vocal apparatus but also that the distance at which long calls are loud
enough for a receiver to be of use to extract individual cues might be less than the
maximum distance at which long calls are audible. The elevation of our playback
speaker (10 m) could also have led to a reduction of the distance versus playbacks
from higher elevation, though male orangutans vocalize mostly between 5 and
15 m high in Tuanan (Lameira et al., unpubl. data).

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A.R. Lameira, S.A. Wich

A potential restriction of our study is that we did not outfit the speaker with a
structure resembling the cheek flanges of a flanged orangutan male. If the flanges are
important to channel long calls in a particular direction, they might reduce distortion
of certain characteristics.
Our results indicate that long calls can indeed serve as long-distance calls from
which receivers can distill individuality, even when some parameters decrease
significantly over distance. Further playbacks with experimentally manipulated long
calls are needed to examine which acoustical parameters are most salient to
orangutans for individual recognition.
Acknowledgments We thank to the Indonesian Institute of Sciences (LIPI) for authorization to carry out
research in Indonesia, the Borneo Orangutan Survival Foundation (BOS) for permission to work at
Mawas, and the Universitas Nasional (UNAS) for acting as a sponsor. The Calouste Gulbenkian
Foundation (Lisbon) and the Leonardo da Vinci Programme (University of Lisbon) financially supported
A. Lameira during data analysis. The Netherlands Organisation for Scientific Research (NWO), the
Leakey Foundation, and the National Geographic Society financially supported S. A. Wich. We thank
Madeleine Hardus, Han de Vries, and Adrian Jaeggi for help during various phases of the project. We
thank Carel van Schaik for providing us the opportunity to conduct research in Tuanan. Two anonymous
reviewers provided valuable suggestions on an earlier version of the manuscript.

References
Brown, C. H. (2003). Ecological and physiological constraints for primate vocal communication. In A. A.
Ghazanfar (Ed.), Primate audition: Ethology and neurobiology (pp. 127149). Boca Raton, FL: CRC
Press.
Cheney, D. L., & Seyfarth, R. M. (1980). Vocal recognition in free-ranging vervet monkeys. Animal
Behaviour, 28, 362367.
Cheney, D. L., & Seyfarth, R. M. (1999). Recognition of other individuals social relationships by female
baboons. Animal Behaviour, 58, 6775.
Davila Ross, M. (2003). The long-calls of wild male orangutans: A phylogenetic approach. PhD thesis,
Tierarzliche Hochschule Hannover, Hannover.
Davila Ross, M., & Geissmann, T. (2007). Call diversity of wild male orangutans: A phylogenetic
approach. American Journal of Primatology, 69, 305324.
Delgado, R. A. (2003). The function of adult male long-calls in wild orangutans (Pongo Pygmeus). PhD
thesis, Duke University, Durham, NC.
Delgado, R. A. (2007). Geographic variation in the long calls of male orangutans (Pongo spp.). Ethology,
113, 487498.
Delgado, R. A., & van Schaik, C. P. (2000). The behavioral ecology and conservation of the orangutan
(Pongo pygmaeus): A tale of two islands. Evolutionary Anthropology, 9, 201218.
Dominy, J., Lucas, P., Osorio, D., & Yamashita, N. (2001). The sensory ecology of primate food
perception. Evolutionary Anthropology, 10, 171186.
Fischer, J. (2004). Emergence of individual recognition in young macaques. Animal Behaviour, 67, 655661.
Galdikas, B. M. F. (1983). The orangutan long-call and snag crashing at Tanjung Puting Reserve.
Primates, 24, 371384.
Hardus, M. E., Lameira, A. R., Singleton, I., Knott, C. D., Morrogh-Bernard, H., Ancrenaz, M., et al.
(in press). The orangutan vocal and sound repertoire: With a focus on geographical variation. In S. A.
Wich, S. Utami-Atmoko, T. Setia, & C. P. van Schaik (Eds.), Orangutans: Geographical variation in
behavioral ecology. Oxford University Press.
Hochberg, Y. (1988). A sharper Bonferroni procedure for multiple tests of significance. Biometrika, 75,
800802.
Hunter, M. L., & Krebs, J. R. (1979). Geographical variation in the song of the great tit (Parus major) in
relation to ecological factors. Journal of Animal Ecology, 48, 759785.
Konishi, M. (1970). The evolution of design features in the coding of species-specificity. American
Zoologist, 10, 6772.

Orangutan Long Call Degradation

625

MacKinnon, J. (1974). The behaviour and ecology of wild orang-utans (Pongo pygmaeus). Animal
Behaviour, 22, 374.
MacKinnon, J. (1979). Reproductive behavior in wild orangutan populations. In D. A. Hamburg, & E. R.
McCown (Eds.), The great apes (pp. 257273). Menlo Park CA: Benjamin/Cummings.
Marten, K., Quine, D., & Marler, P. (1977). Sound transmission and its significance for animal
vocalization. II. Tropical forest habitats. Behavioral Ecology and Sociobiology, 2, 291302.
Mitani, J. C. (1985). Sexual Selection and adult male orangutan long-calls. Animal Behaviour, 33, 272283.
Mitra Setia, T., & van Schaik, C. P. (2007). The response of adult orang-utans to flanged male long calls:
Inferences about their function. Folia Primatologica, 78, 215226.
Morton, E. G. (1975). Ecological sources of selection on avian sounds. American Naturalist, 109, 1734.
Rendall, D., Owren, M. J., Weerts, E., & Hienz, R. D. (2004). Sex differences in the acoustic structure of
vowel-like grunts vocalizations in baboons and their perceptual discrimination by baboon listeners.
Journal of Acoustical Society of America, 115, 411421.
Richards, D., & Wiley, R. (1980). Reverberations and amplitude fluctuations in the propagation of sound
in a forest: Implications for animal communication. The American Naturalist, 115, 381399.
Rijksen, H. D. (1978). A field study on Sumatran orangutans (Pongo pygmeus abelii, Lesson 1827):
Ecology, behaviour and conservation. Veenman: Univeristy of Wageningen.
Schrmann, C. L., & van Hooff, J. A. R. A. M. (1986). Reproductive strategies of the orang-utan: New
data and a reconsideration of existing sociosexual models. International Journal of Primatology, 7,
265287.
Singleton, I., & van Schaik, C. P. (2002). The social organization of a population of Sumatran orang-utans.
Folia Primatologica, 73, 120.
Waser, P., & Brown, C. H. (1986). Habitat acoustics and primate communication. American Journal of
Primatology, 10, 135154.
Waser, P. M., & Waser, M. S. (1977). Experimental studies of primate vocalization: Specializations for
long-distance propagation. Zeitshrift fr Tierpschychologie, 43, 239263.
Wich, S. A., Koski, S., de Vries, H., & van Schaik, C. P. (2003). Individual and contextual variation in
Thomas langur male loud calls. Ethology, 109, 113.
Wiley, R. H. (1991). Associations of song properties with habitats for territorial oscine birds of eastern
North America. The American Naturalist, 138, 973993.
Wiley, R. H., & Richards, D. G. (1978). Physical constraints on acoustic communication in the
atmosphere: Implications for the evolution of animal vocalizations. Behavioral Ecology and
Sociobiology, 3, 6994.