On the Diurnal Advertisement Call Frequency of Hemidactylus Frenatus with Additional Remarks on the Distress Call and Churr Call

2010·3.8 MB·

by  Dieter Gramentz

Journal ofthe Bombay Natural HistorySociety, 106(2), May-Aug 2009 135-141
ON THE DIURNALADVERTISEMENT CALL FREQUENCY
OF HEMIDACTYLUS FRENATUS WITH ADDITIONAL REMARKS
ON THE DISTRESS CALLAND CHURR CALL
DieterGramentz1
'FolderichstraBe7, D-13595 Berlin,Germany. Email: [email protected]
VariousaspectsofthebioacousticbehaviourofHemidactylusfrenatuswerestudiedin November2007 inAluthgama,
Western Province, Sri Lanka. Markedly increased production of advertisement calls was noted about 30 to 50 min
priorto sunsetorabout 70 to 90 min priorto complete darkness; andduring most nights (n=8), peakcalling activity
was observed during dusk from 1750-1830 hrs, between sunset and complete darkness. Advertisement call activity
wasfoundtobemuchreducedduringnightswithprolongedrainincomparisontonightswithoutrain,andthedifference
wasstatistically significant(P<0.05). Minimumnumberofcalls within7 hoursrecordingwas59onarainy nightand
208duringadrynight.Averagenumberofadvertisementcallsonrainynightswas60.5 (SD=2.12;range: 59-62;n=2),
while average number of calls on dry nights was 144.9 (SD=35.7; range: 110-208; n=8). There was statistically
significant0-0.63; P<0.05)correlation between the numberofadvertisementcalls and average airtemperature.The
distress call is a short, relatively high-pitched squeak and its average length was 0.041 sec (SD=0.03; range:
0.013-0.080 sec; n=5). Average maximum sound intensity was 89.7 dB (SD=10.69; range: 79.5-105.8 dB; n=5).
Maximumsoundintensitywasreachedbetween3,967and5,443Hz(x=4,871 Hz;SD=592;n=5). Maximumrecorded
frequencywas 18,636Hz,butmaximumfrequencycanbeaslowas 12.455 Hzwith an averageof 14,835 Hz. Lowest
callfrequenciesrangedfrom 554to 1,199 Hz(x=904Hz; SD=287; n=5).Thesnare-likechurrcall wasstructuredasa
numberof6pulses. Pulselengthsvariedbetween0.006and0.007 sec (x=0.0063 sec; SD=0.005; n=6), andtimegaps
betweenpulseswere0.021 to0.026sec(x=0.023sec;SD=0.003;n=5).Churrcalllength was0.160secandmaximum
sound intensity was 76.4 dB reached at 5,440 Hz. Minimum and maximum frequency was 369 and 15,869 Hz
respectively.
Key words: Hemidactylusfrenatus, bioacoustics, advertisement call, distresscall, churrcall. Sri Lanka
INTRODUCTION
AccordingtoDaniel(1983),thespeciesisperhapsthenoisiest
ofIndiangeckos.Territorialadvertisementcallsaresupposed
For many decades the presence of a voice in geckos tobethemeansforspacingthemselvesouttoclaimareasfor
has been well-known. However, it was not until 1968 and feeding and breeding.
1969whenthefirstanalysisofadvertisementcallsinbarking Despitethewell-studiedstructuresofthedifferentcalls
geckos Ptenopus garrulus and P. kochi respectively, were ofH.frenatus (Marcellini 1974, 1977a) not much is known
,
carried out by Haacke. Since then advertisement calls have onitsdiurnalrhythmicity. Hediger(1934)briefly mentioned
been the subject of research in a number of gecko genera, that H. frenatus not only calls during dusk, but occasionally
e.g., Ptyodactylus (Frankenberg 1973; Werner et al. 1978), also during the day. The species was reported by McCann
Hemidactylus (Marcellini 1974, 1977b; Frenkel 2006), (1940) from Sutgutti, India, to be very vociferous in June
Tarentola (Nettmann and Rykena 1985) and Thecadactylus andcallingfrequentlyatintervalsall night.Anothermention
(Gramentz2007b).Anothergeckocallonwhichbioacoustical ofthe voice ofH.frenatus stems from Poulin etal. (1995),
research isconcentratedisthedistresscall. Distresscall was whichreported"growlcalls”duringaggressiveinteractions.
studied by Frankenberg (1975, 1978), Gramentz and Barts While describing the different calls of H. frenatus,
(2004), Gramentz (2004, 2005b, 2005c) and Barts (2006). Marcellini (1974) did not name them according to the
Brown(1984/85)evennotedanultrasoundcomponentinthe behaviouralcontextinwhichthecallswereused,butinstead
distress call ofmany gecko species. differentiated them by their sound effect and number of
Hemidactylusfrenatusisafamiliarhousegeckospecies syllables emitted (e.g. churr call, single chirp call, multiple
andknowntobe vocally veryactive. The advertisementcall chirpcall).Marcellini(1974)roughlyreportedthatthedistress
of H. frenatus is well-known and they are even called call is very short, < 0.05 sec, and that it begins and ends
“tinktock” or “tschicktschack” (Manthey and Grossmann abruptly.Thedominantfrequencyisapproximately2,000Hz,
1997). The advertisement call of H. garnotii another well with harmonics at 1,000 Hz interval above the dominant
,
known call, is called “tjik tjak” in Malaysia (Steck 1908). frequency. He only published audiospectrograms and these
DIURNALADVERTISEMENT CALL FREQUENCY OF HEMIDACTYLUSFRENATUS
were very much compressed on the time scale and did not Table 1:Airtemperaturesatwhich advertisementcalls
allow apropercall structure analysis. Untilnow the calls of of Hemidactylusfrenatuswere recorded atAluthgama,
Western Province, Sri Lanka
H.frenatus were studied only in subpopulations into which
the geckos were introduced by human activities, such as Date x(°C) SD Range (°C)
Mexico (Marcellini 1974) and Costa Rica (Frenkel 2006). Nov. 08, 2007 25.7 1.08 24.5-27.4
Nov. 09, 2007 27.5 1.00 26.1-29.6
Thepresentstudywillshowaspectsofthespecies’bioacoustic
Nov. 10, 2007 27.7 1.04 26.3-29.9
behaviourin its native environment. Nov. 11, 2007 27.4 1.54 25.0-31.0
Nov. 12, 2007 25.9 1.65 24.4-29.0
MATERIAL AND METHODS Nov. 13, 2007 28.3 1.40 26.3-31.6
Nov. 14, 2007 27.4 2.04 25.0-31.9
Nov. 15, 2007 27.1 1.51 25.4-30.2
Toevaluate overall advertisementcalling activity from Nov. 23, 2007 27.6 1.25 25.9-30.3
one location, Aluthgama, western Sri Lanka (6°25'48.89 N; Nov. 24, 2007 27.3 1.60 25.9-30.9
79°59'54.35E),alladvertisementcallsofH.frenatuswhichcould
beheardwerenoted.Recordingtimewasbetween 1700hrsand Besidesnighttime,H.frenatusproducesadvertisement
2400 hrs. Time ofdusk, sunset and total darkness was noted. calls during the day, but comparatively rarely. Calls ofthis
Additionally the air temperature was recorded at 30 min type were noted duringdaylight atmorning (e.g., 0838 hrs),
intervals startingat 1700hrs andendingat 2400hrs resulting midday (e.g., 1302hrs),andafternoon(e.g., 1514hrs)hours.
in 15 measurements per night. The digital thermometer was It showed that calling activity started about 30 to 50
installedwithathermocoupleataheightof2m.Furthermore, min prior to sunset or about 70 to 90 min prior to complete
weatherandmeteorologicalparametersasclearandovercastsky, darkness. Duringmostnights(n=8)peakcallingactivitywas
rain and thunderstorms were also noted. Recording dates were noted precisely at 1750-1830 hrs (Fig. lb; f to j) between
eightconsecutivenightsfromNovember08toNovember 15,2007 sunsetandcompletedarknessorjustbefore sunset (Fig. le),
and another two consecutive nights on November 23, and oratcompletedarkness(Fig. la).Injusttwocases,thecalling
November 24, 2007. Judged from the various different activity pattern showed a different distribution (Fig. lc, d).
directionsofwhichthecallscouldbeheard,theypossiblycame On all days a sharp increase in calling activity could be
fromabout 10-15 male H. frenatus. observed from about 1700 and 1730 hrs onwards. On two
Additionally five distress calls and a churr call were consecutivenightsofNovember 10and 11,2007 (Fig. lc,d)
recordedandanalysed.Therecordingequipmentisthe same the pattern of calling was different from other nights, but
describedby Gramentz(2005a, c). The soundcardusedwas similaron thesetwonights.Herepeakcallingactivitywasat
Creative Soundblaster Audigy 2 ZS Platinum Pro with a about2115hrsand2130hrsrespectively,i.e.,ashiftofthree
samplerateof44,100Hz, 16bit.Varioussoftwareswereused tothree andahalfhours in comparisontomostothernights.
for sound analysis, such as Avisoft-SASLab, Creative At times akindofdynamicsintheproduction ofcalls
WaveStudioandRaven1.2.Airtemperaturesatwhichthecalls can be heard. These may result in short peaks in calling
wererecordedrangedfrom27.6-30.6°C (Table 1). Distance activity. Example, two males respond to the advertisement
fromthegeckostothemicrophonewhilerecordingchurrand call ofone male, followed shortly by othermales in hearing
distresscalls was 5-10 cm. distance. So, occasionally a fairly large number of calls
Terminology was used as in Gramentz (2003, 2008), (e.g., 8callsin 5 minutes)canbeheardinarathershorttime
however, “churrcall” was adoptedfrom Marcellini (1974). from differentdirections.Aftersome time when mostmales
inthe vicinity have produced one ortworesponsecalls, call
RESULTS frequency is reduced to a lower rate until this kind of
escalation pattern repeats. The result is a rather wavy
Advertisement Cali appearance ofcalling activity during the recording time.
As previously described by Marcellini (1974) a Besidethereactionofreplyingtoanadvertisementcall
repertoire ofthree different call types could be identified in thereisanothersituationwhensuchacallisemitted.Anumber
H.frenatus in Sri Lanka. The production ofthese calls was oftimes I observed that one advertisement call is produced
clearly situation dependent. Directly during an aggressive afteramalegeckosuccessfullychasedawayanintruderfrom
encounter between two males, a short trill-like call can be his territory. The victorious gecko immediately returned to
produced.This“churrcall”isemittedwhenonemalechases histerritory,formedanarchwithitsbody andemittedacall.
anotherinordertodriveitawayfromitsterritory.Threatand Each call is accompanied by a strong exhalation ofairfrom
distress calls are emitted inthe emotional state offear. the lungs thatcan be easily observed from the side.
136 J. Bombay Nat. Hist. Soc., 106 (2), May-Aug 2009
a
9 08NOV2007
d
m535ms;;m;;;m;jgiggSn»55mS!jmsSSm!!SmES
KS$;s£sjsts;sts:gjs;gjs;gjs
Fig. 1 a-j: Frequencyofadvertisementcalls of Hemidactylus frenatusfrom Sri Lanka during different nights
Left dotted line marksthe timeof sunset, right dotted line marks pointofcomplete darkness
1 Bombay Nat. Hist. Soc., 106 (2), May-Aug 2009 137
DIURNALADVERTISEMENTCALL FREQUENCY OF HEMIDACTYLUSFRENATUS
_U5 -|
"0ra 200 -' •
1
:
E
0
1 150 - . .
0 •
>
~o •
TO
1°4- 100 - • •
0
E
^ r—
50 -1 , , , , , , , , , > , ,
25.6 25.8 26.0 26.2 264 26.6 26.8 27.0 27.2 27,4 27.6 27.8 28.0 28.2 28.4
Airtemperature (°C)
Fig.2:VariationoftheadvertisementcallactivityofHemidactylus Fig. 3: Relationship between advertisement call frequency of
frenatusfrom Sri Lanka during rainyand dry nights Hemidactylusfrenatusand airtemperatureduring 10 nights
A lPillPlii'lll
1
0.15 0.2 0.25 0i3 0.35 s
Fig.5:Audiospectrogramofadistresscallofamale Hemidactylus
Fig. 4: Oscillogram of a distress call of a male Hemidactylus frenatus from Sri Lanka. Length of the call is 0.080 sec.
frenatusfrom Sri Lanka. Length ofthe call is 0.083 sec The harmonics are at the frequencies 3,414 Hz, 5,351 Hz,
7,012Hz,8,765Hz, 10,425Hz, 12,086Hz, 13,898Hzand15,592Hz
Advertisement call activity is very much reduced (Figs4 and 5). The average length ofthe recorded calls was
during nights with prolonged rain in comparison to nights 0.041 sec (SD=0.03; range: 0.013-0.080 sec; n=5).
without rain. Minimum calling activity was 59 calls on Maximum sound intensity varied between 79.5 and
November08, duringthe seven hoursrecording,ofwhichit 105.8 dB. On an average maximum sound intensity in the
was raining for 1 hr 33 min. On November 12, it rained for five recorded distress calls was 89.7 dB (SD=10.69).
1 hr 8 min, and 62 calls were recorded. Maximum number Maximum recorded frequency ranged from 12,455-
of calls (208) was recorded on November 11, a dry night. 18,636 Hz, with an average of 14,835 Hz. Lowest call
Average numberofadvertisement calls during rainy nights frequencies varied between 554 and 1,199 Hz having an
was60.5 (SD=2.12; range: 59-62; n=2).Average numberof averageof904Hz(SD=287;n=5).Theaverageofmaximum
callsduringdrynightswas 144.9(SD=35.7;range: 110-208; frequency in the five distress calls was 14,835 Hz. Despite
n=8), and there is a statistically significant difference the wide frequency span ofabout 11-18 kHz in the recorded
(P<0.05; r=3.197, t-test)betweenthemeansofadvertisement distresscallsthe spanatwhich themaximumsoundintensity
calls on rainy and dry nights. Overall average was 128 calls is produced covers a rather small range of about 1.5 kHz.
between 1700-2400 hrs in 10 nights. Maximum sound intensity was found to be between 3,967
Call activity seems to be influenced by the weather, and5,443Hz(Figs6and7).Theaveragefrequencyatwhich
sincetherewasincreaseincallingactivityondrynightswith maximumsoundintensitywasnotedwas4,871 Hz(SD=592;
a sharp drop when these were interrupted by a rainy night n=5).
(Fig. 2). Furthermore, call activity was positively related to The distress call of H. frenatus shows rather
air temperature. There was a modest but statistically similar intervals between harmonics. The average interval
significant (/=0.63, P<0.05) correlation between the total between harmonics of the distress call shown in Fig. 5
number of advertisement calls and average air temperature was 1,740 Hz (SD=104.2; n=7). The interval between
during the night (Fig. 3). harmonics ranged from 1,660 Hz to 1,937 Hz. Frequency
of the lowest and at the same time strongest harmonic
Distress Call was 3,414 Hz. The highest harmonic had a frequency of
The distress call is a very short high pitched sound 15,592 Hz.
138 J. Bombay Nat. Hist. Soc., 106 (2), May-Aug 2009
DIURNALADVERTISEMENT CALL FREQUENCY OF HEMIDACTYLUSFRENATUS
Fig. 6: Three-dimensional logarithmic image ofa distress call of Fig. 7: Three-dimensional logarithmic image ofadistresscall of
individual-a Hemidactylusfrenatus(male) individual-b Hemidactylusfrenatus(male)
0.4 0.5 0.6 0.7 0.8 s
Fig.8:OscillogramofachurrcallofamaleHemidactylusfrenatus Fig. 9: Audiospectrogram of the same churr call of a male
from Sri Lanka. Length ofthecall is 0.160 sec Hemidactylusfrenatus
ChurrCall no marked peak early after this initial calling activity.
The chuiTcall consists ofsix strongerdistinguishable Obviously there is ageographical difference in peakcalling
pulses, which can be identified in the oscillogram and activitybetweenthetwolocations.AlsoinPtenopusgarrulus
audiospectrogram(Figs8and9).The lengthofapulsevaries peak calling activity was noted at sunset when darkness
between0.006and0.007sec(x=0.0063sec;SD=0.005;n=6). increases (Brain 1962). According to Loveridge (1947),
Thetimegapbetweenthesepulsesvariedbetween0.021 and P. garrulus calls during the short period of twilight. The
0.026 sec (x=0.023 sec; SD=0.003; n=5). phenomenonthatH.frenatusmayshowdifferentpeakcalling
The single churr call had a length of 0.160 sec. activities requires further investigation from other
Maximum sound intensity was found to be 76.4 dB, which geographically different locations.
was less than in the weakest distress call. Maximum sound Ihavetheimpression,althoughthisisnotyetconfirmed
intensity was howeverreached at 5,440 Hzwithin the range bydirectobservationofacertainindividual,thatamalemay
ofthe recorded distress calls. The lowest calling frequency givehisfirstadvertisementcall justaboutthesametimewhen
wasmeasuredinthechurrcallwithjust369Hz,butmaximum itstartsactivity forthenight.Also,Marcellini(1974)reported
frequency of 15,869 Hz was comparable to the range ofthe that after emergence from their diurnal retreats, geckos
distress call. commonly called before moving to their feeding areas. It is
obviously of major importance for the geckos to announce
DISCUSSION territorialitypriortothe start ofnocturnalactivity.Asinthis
study, Manthey and Grossmann (1997) noted that calls of
Marcellini (1974) recorded advertisement calls H. frenatus can also be heard during the whole day, and
(multiple chirp calls in his terminology) ofH.frenatus per Marcellini (1974) wrotethatfewcallsoccurduringdaylight
hourinMexicoduringfiveconsecutivenightsandcondensed hours. As in this study, also Frenkel (2006) found that call
theresultsintoonegraph. Healsonotedanincreaseincalling activity ofH.frenatusstudiedinPuntaMorales, CostaRica,
activityathisstartingpointat 1800hrs. However,heobserved was positively correlatedto airtemperature at night.
asteadyincreaseincallingactivityinthegeckosfromMexico Advertisementcallswhichareformedbyalargenumber
withapeakatabout0330hrsinthe night.Theearly increase ofratheridenticalsyllablesareknownfromotherHemidactylus
incallingactivity aroundsunsetresemblesthefindingsfrom species: H. angulatus (Gramentz 2005d), H. mabouia
Sri Lanka, but contrary to the geckos from there, there was (Gramentz2003;Regalado2003),//.platycephalus(Gramentz
J. Bombay Nat. Hist. Soc., 106 (2), May-Aug 2009 139
.
DIURNALADVERTISEMENTCALL FREQUENCYOF HEMIDACTYLUSFRENATUS
2005a)and//, turcicus(Marcellini 1977a;Frankenberg 1982). inakindoftail inwhichevensinglepulsescanbeidentified.
Furthermore, this rather stereotyped territorial call is known In comparison, a distress call actually having an abrupt
from other genera as Phyllodactylus (Marcellini 1977b), beginning and ending is for example produced by
Ptenopus(Haacke 1968, 1969;Gramentz2008),Ptyodactylus Haemodracon riebeckii (Gramentz 2005b). There are also
(Frankenberg 1973, 1974), Tarentola (Nettmann and Rykena some differences in the overall frequencies and intervals in
1985) and Thecadactylus (Gramentz 2007b). Multiple chirp thedistresscallsrecordedat Sri Lankaandthe datareported
calls can, however, also have a submissive function as in by Marcellini (1974) from Mexico. He mentioned as the
Cosymbotusplatyunis(Gramentz2007a). dominant frequency 2,000 Hz with harmonics at 1,000 Hz
From H. angulatus and H. platycephalus another call intervals. InSriLanka,thisfrequencywashigher(3,414Hz)
consistingofalargenumberofsyllablesisknown(Gramentz and the interval between harmonics averaged 1,740 Hz.
2005a,d).Thiscontactcallhasaratherweaksoundintensity According to Marcellini (1974) the churr call is an
andisdisplayedbythemaleinclosemale-femaleinteraction. infrequently heard vocalization and he recorded twice of
Itwouldbeveryinterestingtoknowwhetherthistypeofcall which both were less than 0.2 sec long. I also recorded this
is also a part ofthe repertoire ofH.frenatus. kind ofshort call duration, having a length of0.16 sec. He
Marcellini (1974)wrotethatthedistresscall(hissingle furtherobservedthatthechurrcallwasaudiblefromadistance
chirp call) is less than 0.05 sec long. In the present study, it of 35 m. As this type of call was the weakest recorded at
showedthatthis typeofcall is indeed very shortinduration. Sri Lanka, it is likely that, similarly as in the distress call, a
In fact, the shortest calls were just 0.013 and 0.016 sec high variation of sound intensity exists. The growl calls
long, however, two distress calls had lengths of 0.060 and reported by Poulin etal. (1995) are most probably identical
0.080 sec. He also noted that some calls can only be heard to the churrcalls first described by Marcellini (1974).
fromafewmetresawaywhileothersareclearlyaudiblefrom Only maleswerefoundtoemitchurranddistresscalls
10m.This isreflected in the very different sound intensities at Sri Lanka. Marcellini (1974) reported that only males
of79.5 to 105.8dB. Likehim, Icannotexplainthereasonfor emitted churrcalls and this isconsistentwith thefindingsof
thesevariationsinsoundintensityinthedistresscall. Distress H. frenatus at Sri Lanka.
calls are already known from othergecko species to vary in Marcellini’s (1974) sound analysis equipment seems
length.InStenodactylusstenurus threedifferentdistresscalls to be restricted in detecting frequencies above 8,500 Hz as
,
were noted varying mainly in length, but also in sound his graphs of audiospectrograms showed maximum values
intensity (Gramentz 2004). of6or8kHzonthey-axis.Therefore,theimpressionappears
Possibly due to the equipment used by Marcellini that the call frequencies reach their full capacity within this
(1974) he got the impression of distress calls abruptly range. This is, however, not the case. Both the churr and
beginning and ending. However, as shown in Fig. 3 the distress call reach frequencies above 15 and 18 kHz
intensityincreasestoamaximumafter0.22sec.Thecallends respectively (Figs 5, 6, 7 and 9).
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