Comparison of anuran acoustic communities of two habitat types in ...

Anuran acoustic communities in Sabah, Malaysia 

SALAMANDRA 

43 

3 

129-138 

Rheinbach, 20 August 2007 ISSN 0036-3375 

Comparison of anuran acoustic communities of two 

habitat types in the Danum Valley Conservation Area, 

Sabah, Malaysia 

Doris Preininger, Markus Böckle & Walter Hödl 

Abstract. We compared advertisement calls of frog assemblages in two different habitats, (i) an open area 

along a dirt road with ponds and secondary vegetation; (ii) a fast flowing stream in primary forest. Eleven 

frog species were recorded and significant differences in the dominant call frequencies between the two 

observed frog communities were discovered. Stream-breeding species produce higher frequencies than 

species occurring in the roadside habitat. Noisy habitats have an influence on dominant frequency and 

demand acoustic adaptations to increase the signal-to-noise ratio. Selective logging represents a major 

threat to stream-breeding anurans in Sabah. Pollution of clear water threatens the stream-dependent 

herpetofauna. 

Keywords. Amphibia, Anura, conservation, vocalisation, Malaysia. 

Introduction 

Selective logging has been the main cause of 

disturbance to tropical forests in Southeast 

Asia (Marsh & Greer 1992, Willott et al. 

2000). Concession-based timber extraction 

and plantation establishment result in highly 

fragmented forests (McMorrow & Talip 

2001, Curran et al. 2004) and increase 

river sediment yields (Douglas et al. 1993). 

The extent to which biodiversity is altered 

in selectively logged forest is an important 

conservation issue. Fragmentation, degradation 

and conversion of rainforests affect the 

tropical herpetofauna (Alcala et al. 2004). 

About half the frog species in Southeast Asia 

are restricted to riparian habitats and develop 

in streams (Inger 1969, Zimmerman & 

Simberloff 1996). Most anuran stream-side 

communities in Borneo are known to breed 

in clear, turbulent water and are absent in 

streams with silt bottoms and lacking riffles 

and torrents (Inger & Voris 1993). Several 

anuran community studies have focused on 

environmental conditions such as vegetation, 

elevation, rainfall and microhabitat (Inger 

1969, Duellman 1978, Inger & Voris 

1993, Gillespie et al. 2004). Apart from behavioural 

and morphological adaptations to 

the environment, frog assemblages also show 

acoustic differences. A prime factor separating 

species within anuran assemblages 

is vocalization (Duellman & Trueb 1986). 

Acoustic partitioning through temporal and 

spectral features minimizes competition and 

allows conspecific females to recognize individual 

signalling males (Hödl 1977, Garcia- 

Rutledge & Narins 2001, Gerhardt & 

Huber 2002). The dominant frequency of a 

frog’s call is partially constrained by its body 

size (Kime et al. 2000). Snout-vent length and 

dominant frequency are negatively correlated 

(Littlejohn 1977, Duellman & Pyles 1983, 

Ryan & Brenowitz 1985). Sounds produced 

by waterfalls and fast-flowing streams contribute 

to the ambient noise level. Distinct, 

acoustic habitat properties (“melotops”) demand 

different adaptations on animal vocalizations. 

In this study, species composition 

was determined for two habitats within the 

Danum Valley Conservation Area (DVCA), 

a fast flowing stream in the primary forest 

© 2007 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT) 

http://www.salamandra-journal.com 

129

Doris Preininger et al. 

and a roadside habitat with secondary vegetation, 

and spectral and temporal features of 

the acoustic signals were compared. 

Material and methods 

Study area 

From 10 January to13 March 2006, we studied 

frog communities in DVCA, Sabah, Malaysia. 

DVCA has an area of 43,800 ha of lowland 

tropical rainforest and is located in the upper 

catchment area of the Segama River. The Danum 

Valley Field Centre (DVFC) is located 

on the western edge of the DVCA (4°57´40” 

N 117°48´00” E; all but 9% of the area is below 

760 m above sea level) from where a 2.8 km 

long trail leads south to the Tembaling waterfall. 

The rainy season in Sabah extends from 

December to March (Northeast Monsoon) 

with rainfall peak during February (monthly 

mean 788 mm). Annual precipitation is 

near 3000 mm with year-to-year fluctuation 

of about 100 mm. Mean annual temperature 

is 26.7°C with mean maximum 30.9°C and 

mean minimum 22.5°C. Daily temperature 

fluctuations are noticeably more pronounced 

than the year-to-year fluctuations. In the period 

from January to March the highest recorded 

temperature was 32.5°C and the lowest 

was 20.3°C; mean relative humidity at 

the DVFC was 97.0% at 8:00h and 89.7% at 

14:00h. Daily rainfall and temperature data 

from January to March 2006 derived from 

the Danum Valley weather station, which is 

maintained by the staff of the Royal Society 

SE Asia Rain Forest Research Programme. 

Sampling methods 

We focused on two macrohabitat types. One 

is pristine, the other disturbed. The Tembaling 

River within the primary dipterocarp forest 

at about 900 m elevation is a fast flowing 

freshwater stream with rapids, waterfalls and 

no siltation. The selected disturbed habitat 

was along a dirt road in the DVFC with secondary 

vegetation, ephemeral roadside pools 

filling seasonally with water and varying degrees 

of human disturbance. In both habitats 

1000 m long and y-shaped transects were 

established and marked with coloured tape. 

We performed visual and acoustic encounter 

surveys. The search areas also included adjacent 

riverbanks or vegetation up to a distance 

of 3 m to the water- or road-edge. Sampling 

was performed independent of prevailing 

weather conditions. Each transect was sampled 

at least 20 times by two observers during 

the day and during nocturnal censuses. Repeated 

controls of the same transect on consecutive 

days were avoided to ensure independence 

of samples. To determine all calling 

species, a 24-hour time period was sampled 

four times during one month along both 

transects. After locating a vocalizing male, 

call recordings were made from distances of 

1 to 5 m using directional and surround microphones 

(Sennheiser Me 66, AKG D 190 E) 

and DAT- recorders (Sony DAT-Rec. TCD- 

D8). Microhabitat temperature and humidity 

were measured with a digital thermohygrometer 

(Testo 610 GM) before each recording. 

Frogs were located at their resting sites, 

and if possible, captured and placed in plastic 

bags. Snout-vent length and snout-urostyle 

length were measured using a Wiha calliper 

(± 0.1mm). To differentiate between individuals 

we photographed and compared dorsal 

colour patterns. All individuals were released 

after taking their measurements. 

Data analysis 

We digitized and analysed the recorded calls 

with the sound-analysis software Raven 1.2.1 

at a sampling frequency of 44.1 Hz and with 

a mono 16 bit PCM Input and a 10 Hz update 

rate at normal speed. Power spectra, sonograms 

and oscillograms of 1 to 5 advertisement 

calls were analyzed for each frog and 

the average dominant frequency, minimum 

and maximum frequency, call duration, 

note duration, repetition rate and number 

130


of notes, when applicable were calculated 

for each individual. Spectrogram views were 

produced with a Hann-filter with a sample 

size of 512 and a 3-dB filter bandwidth of 124 

Hz. The correlation between body size (SVL) 

and dominant frequency was calculated with 

a Spearman non-parametric correlation (p ≤ 

0.05). ANOVA (p ≤ 0.05) was used to test for 

statistical differences, in SVL (mm) and dominant 

frequency (Hz) of advertisement calls, 

among frog assemblages in two habitats. The 

influence of SVL and habitat on the dominant 

frequency of the frog species was calculated 

by an ANCOVA (p ≤ 0.05). Since the sample 

size for some anuran species tested was N = 1, 

the influence of SVL on dominant frequency 

was calculated twice: once corrected according 

to number of individuals and once without 

correction. All statistical tests were produced 

with SPSS for Mac 11.0.4. 

(Fig. 5), Polypedates otilophus (Fig. 6), and 

Bufo asper (Fig. 7), Meristogenys orphnocnemis 

(Fig. 8), Microhyla sp. (Fig. 9), Staurois 

natator (Fig. 10), Staurois latopalmatus (Fig. 

11), respectively. No calls could be recorded 

for species without frequency data (Tab. 1). A 

total of 213 calls were analyzed. The dominant 

call frequencies of the anuran species are not 

dependent on the SVL of the vocalizing individuals 

(N = 11; c = -0.555; p ≤ 0.077; r 2 = 

0.30) (Fig. 13). Meristogenys orphnocnemis, a 

species found in the stream habitat produced 

the highest calling frequency (7205.0 Hz). 

Frog species from the torrent are above the 

Results 

During the sampling period 20 frog species 

were found. Eleven frog species were recorded, 

six along the disturbed area and five within 

the pristine habitat: Fejervarya nicobariensis 

(Fig. 1), Fejervarya limnocharis (Fig. 2), 

Rhacophorus dulitensis (Fig. 3), Polypedates 

leucomystax (Fig. 4), Polypedates macrotis 

Fig. 2. Spectrogram (frequency [kHz]) and waveform 

(rel. amplitude [kUnit] = proportional to 

sound pressure of the recording) of the advertisement 

call of Fejervarya limnocharis. Temperature 

during recording 24.6 °C. 




call and non-collected specimen of Fejervarya 

nicobariensis. Temperature during recording 

25.4 °C. 




call and non-collected specimen of Rhacophorus 

dulitensis. Temperature during recording 26.3 

°C. 

131





call and non-collected specimen of Polypedates 

leucomystax. Temperature during recording 

25.0 °C. 





otilophus. Temperature during recording 

25.4 °C. 





macrotis. Temperature during recording 

24.9 °C. 

calculated regression except Bufo asper and 

Microhyla sp. All frog species in the roadside 

habitat are below the calculated regression. 

The largest frog species was Polypedates 

otilophus with a dominant call frequency of 

1,033.6 Hz. No frog species with a dominant 

call frequency below 1,000 Hz was recorded. 

Frequencies of advertisement calls of anuran 

species found in the stream transect are higher 

than call frequencies recorded of species 

occurring in the roadside habitat (Fig. 14) (N 




call and non-collected specimen of Bufo 

asper. Temperature during recording 24.6 °C. 

= 11; df = 1; F = 7.315, p = 0.024). Dominant 

frequency is not dependent on the SVL (N = 

11; df = 1; F = 3.822, p = 0.082) but the influence 

of the habitat is significant (N = 11; df = 

1; F = 7.031, p = 0.029). If the number of analyzed 

individuals is included, the dependency 

of the dominant frequency, the influence 

of SVL (N = 11; df = 1; F = 0.519, p = 0.049) 

and the influence of the habitat (N = 11; df = 

1; F = 11.435, p = 0.010) become more significant. 

132





call and non-collected specimen of Meristogenys 

orphnocnemis. Temperature during recording 

24.8 °C. 




call and non-collected specimen of Staurois 

natator. Temperature during recording 25.3 °C. 




call and non-collected specimen of Microhyla 

sp. Temperature during recording 25.3 °C. 

Discussion 




call and non-collected specimen of Staurois 

latopalmatus. Temperature during recording 25.2 

°C. 

The comparison of two local frog assemblages 

in different habitats shows that advertisement 

calls of frog species living in noisy 

environments constrain higher frequencies. 

Fast flowing streams and waterfalls produce 

low-frequency-dominated noise which may 

present a selective force for stream-breeding 

species (Hödl & Amézquita 2001, Feng et 

al. 2006). The high-frequency calls exceed 

the constant ambient noise level. Spectrograms 

of streams show high sound pressure 

levels in low frequencies and lower sound 

pressure levels in frequencies above 3,000 

Hz (Hödl & Amézquita 2001, Feng et al. 

2006, authors’ unpublished data). Environmental 

constraints should have an effect on 

the evolution of signals (Narins & Zelick 

1988). Selection favors signals that minimize 

the effects of background noise and interfering 

signals from other species (Endler 

1993). Riparian frog species are acoustically 

well adapted to their environment and show 

133


a 

b 

c 

d 

e 

f 

g 

h 

Fig. 12. a) Fejervarya nicobariensis, b) Fejervarya limnocharis, c) Rhacophorus dulitensis, d) Polypedates 

leucomystax, e) Polypedates macrotis, f) Polypedates otilophus, g) Bufo asper, h) Meristogenys orphnocnemis, 

i) Microhyla sp., j) Staurois natator, k) Staurois latopalmatus. Photos: M. Böckle, W. Hödl, T. 

Reis. 

134


i 

j 

k 

strong habitat affiliations. With the exception 

of Bufo asper the mean dominant frequency 

was above 3.9 kHz for frogs calling in the undisturbed 

habitat. A possible explanation for 

the call frequency in B. asper could be that it 

prefers stream banks, with small riffles that 

produced less background noise. Additionally, 

dominant frequency is determined by 

SVL. Larger frogs call with lower frequencies 

than smaller frogs (Littlejohn 1977, Kime 

et al. 2000). The sampling size was probably 

too small to produce the dependency of 

SVL on dominant frequency. Only weighted 

data were able to show the influence. All species 

found in the disturbed roadside habitat 

call with lower frequencies and less constant 

noise is emitted by the surrounding environment. 

This gives rise to the question why a 

high proportion of species breeds close to 

fast flowing streams (Inger 1969). Zimmerman 

& Simberloff (1996) showed that reproductive 

mode and developmental habitat 

are strongly associated with phylogeny. The 

same limited set of ecological and reproductive 

traits is shared by ranids, pelopatids and 

rhacophorids. Environmental conditions and 

interspecific interactions may have led to adaptations, 

but local selection on species can 

not be counted independent when closely related 

species occur in the same community. 

Phylogeny was not included in our analysis 

and a certain bias may result from the fact 

that two species of the genus Staurois are 

present at the stream habitat. Advertisement 

calls are important determinants of reproductive 

success and could have evolved in a 

common ancestor. However, we were able 

to record five species of the family of ranids, 

three at the stream (Staurois latopalmatus, 

S. natator and Meristogenys orphnocnemis) 

and two along the roadside (Fejervarya nicobariensis 

and F. limnocharis). All three ranid 

species recorded along the stream produce 

advertisement calls in which the dominant 

frequencies are above the calculated linear 

regression line (Fig. 13). Although morphological 

and phylogenetic limitations constrain 

frog vocalizations, selective pressures 

such as environmental factors have resulted 

in a diversity of advertisement calls (Duellman 

& Trueb 1986). Acoustic partitioning 

and increasing the signal-to-noise ratio is a 

prerequisite for frogs living along torrents 

and streams. The influence of the habitat on 

advertisement calls should emphasize the 

importance of microhabitat composition on 

frog assemblages. Most recorded species are 

threatened by habitat loss (Tab. 1). Deforestation 

and subsequent siltation of streams and 

rivers are the major threats to most streambreeding 

species in Sabah. Out of twelve species 

found along the stream, three are classified 

as “near threatened” according to the 

135


Fig. 13. Influence of SVL on dominant frequency of the species advertisement call (y = -55.49x + 

5472.71). Frog species of the stream transect are marked with a square, those from the roadside habitat 

with a circle. 

Fig. 14. Dominant call frequencies of the anuran 

species from two different habitat types. Species 

along the stream call with higher frequencies than 

frog species in the roadside habitat. 

IUCN Red List status (Tab. 1). Inger & Voris 

(1993) found that a stream with a silt bottom 

completely lacked all the species known to 

breed along clear and fast flowing streams. 

Selective logging changes the water chemistry 

considerably in nearby streams. Sediment 

yields of streams are 18 times higher within 

the next five months in selectively logged areas 

(Douglas et al. 1992, 1993). The increase 

in sedimentation levels has a clear effect on 

larval anurans and on reproduction of several 

stream-breeding species. Frog assemblages 

as those found at the Tembaling River show 

the importance of conservation areas like 

Danum Valley. On the other hand more microhabitats 

are created through reduced impact 

logging which attracts frog species that 

are normally not found in primary forests 

(e.g., Polypedates macrotis, Fejervarya nicobariensis 

and Rhacophorus pardalis) (Wong 

in press). We believe that disturbed areas 

such as those around the Danum Valley Field 

Center and adjacent roads contain suitable 

ponds and breeding habitats for disturbancetolerant 

breeders, but the large destruction 

and fragmentation of primary forest poses a 

136


Tab. 1. List of anuran species found including weight, snout-vent length (SVL), dominant call frequency 

(DF) and IUCN Red List status. Habitat: 1 = stream, 2 = road edge (numbers in parentheses are the absolute 

number of individuals measured); weight [g]; SVL [mm]; dominant frequency, DF [Hz] (number 

of individuals/number of analyzed calls); major threat: HL = habitat loss, P = pollution of streams and 

rivers, ID = infrastructure development, N = none; Red List status according to IUCN et al. (2006). 

species habitat SVL DF major threat IUCN Red List 

status 

Bufo asper 1 76.2 (1) 1,031.2 (1/9) HL/P least concern 

Meristogenys orphnocnemis 1 30.2 (4) 7,205.0 (1/6) HL/P least concern 

Microhyla sp. 1 14.9 (5) 3,969.3 (4/25) - - 

Staurois natator 1 33.9 (4) 4,746.0 (3/19) HL/ID least concern 

Staurois latopalmatus 1 47.7 (11) 5,149.4 (5/20) HL/P least concern 

Ansonia longidigita 1 - - HL/P near threatened 

Ansonia spinulifer 1 3.2 (1) - HL/P near threatened 

Leptobrachium abbotti 1 56.5 (5) - HL least concern 

Limnonectes leporinus 1 106.8 (1) - HL least concern 

Rana picturata 1 39.8 (2) - HL least concern 

Pedostibes rugosus 1 - - HL/P near threatened 

Chaperina fusca 1 18.8 (1) - HL least concern 

Chaperina fusca 2 - - HL least concern 

Polypedates leucomystax 2 30.0 (7) 2,056.7 (4/35) N least concern 

Polypedates macrotis 2 57.0 (5) 1,388.9 (1/3) N least concern 

Polypedates otilophus 2 81.1 (9) 1,033.6 (1/2) HL least concern 

Fejervarya limnocharis 2 33.4 (3) 1,274.0 (3/3) - least concern 

Fejervarya nicobariensis 2 41.2 (6) 3,169.7 (5/44) HL/N least concern 

Rhacophorus dulitensis 2 46.6 (4) 1,868.6 (5/47) HL near threatened 

Rhacophorus pardalis 2 43.5 (6) - HL least concern 

Occidozyga laevis 2 31.0 (10) - N least concern 

major threat to the characteristic herpetofauna 

of Sabah. 

Acknowledgements 

We thank the Danum Valley Management Committee, 

the Economic Planing Unit, the Sabah 

Wildlife Department and the Universiti of Malaysia 

Sabah for giving us the opportunity to work 

in the Danum Valley Conservation Area. Special 

thanks go to the Royal Society (RS) and Glen 

Reynolds for their support and commitment to 

make this project possible. Julia Felling and 

Karoline Schmidt assisted during our fieldwork 

and Markus Gmeiner did the photo editing. We 

are very thankful for the statistical help of Adolfo 

Amézquita. Anna Wong acted as our local collaborator. 

Financial support was given by the University 

of Vienna (Brief Scientific Stays Abroad) 

and the DGHT (Wilhelm-Peters-Fonds). The 

Department of Evolutionary Biology (University 

of Vienna) provided the necessary field equipment. 

Last but not least we thank the DVCA and 

RS staff for keeping our spirits up. 

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Authors’ addresses: Doris Preininger, Markus Böckle, Walter Hödl, Department for Evolutionary 

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