Sound Production in the Anemonefishes, Amphiprion clarkii and A. frenatus (Pomacentridae), in Captivity

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1 Japanese Journal Ichthyology Vol.35, No Sound Production nemonefishes, mphiprion clarkii frenatus (Pomacentridae), aptivity Kuei-hiu hen H-Kiu Mok (Received pril 13, 1987) Pomacentrids are well known for ir terririal acoustic behaviours. number papers have been published on acoustic behaviour tlantic species (e.g., Myrberg et al., 1978; Spanier, 1979; Myrberg Spires, 1980; Myrberg Riggio, 1985). Study on sound production anemonefishes mphiprion can be traced back as early as 1930 when Verwey reported that mphiprion akallopisos polymnus emit sounds. Schneider (1964) documented threateng, fightg, shakg sounds mphiprion clarkii, polymnus, frenatus, percula, bcctus. However, no detail data on acoustic parameters latter three species were given his paper. llen (1972) described acoustic behaviour chrysopterus perideraion by givg duration, frequency range repetition pulses se sounds. coustic data on remag 26 mphiprion species are far from complete. The present note gives detail acoustic characteristics clarkii frenatus proposals for producg mechanisms ir sounds. Materials methods coustic recordgs were made laborary. tal 16 live specimens mphiprion clarkii 9 frenatus were kept aquaria (90 ~45 45cm3) with crashed corals as substrate ~ water temperature was mataed around 25 Ž. 12:12hrs light-dark period was mataed. The body length specimens ranged from cm SL clarkii cm SL frenatus. Three five dividuals each species were mataed an aquarium for each recordg. Recordg time was romly selected durg day. miature hydrophone (riiel Kjaer type 8103) suspended aquarium was connected a recordg charge amplifier (riiel Kjaer type 2651) cassette recorder (Sony WM D6). fter sounds were recorded, y were analyzed by a digital sonagraph (Kay 7800) a high resolution signal analyzer (riiel Kjaer 2033) which was attached a X-Y recorder (Hewlett Packard Model 7015) for data output. Results Quantitative data on physical parameters mphiprion clarkii frenatus sounds are listed Table 1. The pop chirp sounds are distguishable by ir domant frequency ranges; pops se two species had a wider frequency range than chirps (Figs. 1-4). Spectral distribution also differed between se sounds; sound energy was peaked at about Hz chirps while energy peak pop was not proment (Figs. 2, 4). Species differentiations frequency range spectral distction are significant (Figs. 1-4). In both species pop sound was predomately produced sgly although it could be emitted a set two (Figs. Table 1. Physical parameters pop chirp sounds mphiprion clarkii frenatus.

2 hen Mok: Fig. 1. Sound Production Sonagrams sgle double pop sounds produced natus (, D). Fig. 2. nemonefishes D by mphiprion clarkii (, ) fre- Spectrograms sgle pop sound produced by mphiprion clarkii () frenatus (). spectra: background noise. 1, 5, 6). This sound was ten delivered immediately prior a chirp (Fig. 7) which was composed a series pulses (or clicks; Figs. 3, 8). ccordg its temporal characteristics, chirp is considered an equivalent sgal llen's "click" chrysopterus perideraion (llen, 1972). The composg series ten cluded 1 7 pulses frenatus (with only one series cludg 22 pulses; Fig. 9) 1 17 pulses clarkii (Fig. 9). frenatus more ten emitted chirps with fewer pulses than clarkii did (Fig. 9, ). otm The chirp sounds se two species are different; chirps frenatus have a lower repetition rate than those clarkii (Fig. 3, ). Durg an agonistic confrontation, appearance pops chirps was accomplished with a slight vertical motion domatg contestant's lower jaw. Such motion which is apparently associated with sound production has not been observed subordate contestants (or loser contest). Therefore, it can be concluded tentatively that subdomance hibits 91

3 魚類 学 J. Ichthyol. 35(1), 1988 Fig. 3. production Sonagrams chirp sound produced by mphiprion clarkii (, ) frenatus (). se ccordg more active The former sounds latter (18%) 3; did so from chirp range it a higher (Fig. atus) have shorter is a longer series is crement vocal fourth pulses teeth se vocalization chirp non- bones may sound examed number biacule- number its Premnus teeth Teeth among teeth while active cerapharyngo- frenatus, 11-), The present or published data dicate that mphiprion clarkii, frenatus, akallopisos, polymnus, chrysopterus, percula, bicctus are vocal species while ephippium, ocellaris, Premts biaculeatus are non-vocal ones (Verwey, 1930; Eibl-Eibesfeldt, 1960; Schneider, 1964; Mariscal, 1972; llen, 1972; hen, 1987). ccordg llen (1972), perideraion is a vocal species. However, we rarely found this species producg sounds laborary condition. These specific differences lead followg ferences. s traspecific terspecific teractions evitably take place se species, vocal signal is not a compulsory element agonistic display anemonefishes. Our data show that vocal species are also differentiated ir vocal activity wards conspecifics. s vocalization is probably associated account The Discussion pharyngo- may cane fewer highest fre- lack (pers. teeth E). more sounds. ocellaris 11D, pop swimbladder clarkii, E, which frenatus, wide gratg 10-, (i.e., 10D, cictrkii, long duration, second more (i.e., Figs. species In frenatus, second se carry vocal ant with properties short species its while encounters per cerabranchial plate perideraion; 343 pulses that branchials This sound acoustic believed branchial chirp (65%) non-harmonic) production has out 20 attached is branchial (Figs. Unlike (i.e., muscle vocal 62 was emission. pop chirp sounds quency for sound encounters up clarkii 1). Judgg obs.), 112 emit rate sonic P<0.001). could Table frenatus (ƒô2=89.77; clarkii produced out only repetition observations, species 73 produce sound (or shakg sound) by its submissive display which was a contuous lateral shakg movement with belly facg domant recipient. Sound energy was restricted a narrow, low frequency b around 200Hz (Fig. 12). Its duration was about 65 msec (Fig. 12). The shakg sound was detectable only durg a vigorous shake a large subordate's body. Its low frequency narrow frequency range suggest that it is a hydrodynamic sound. sounds. our than than species. be relevant characteristics. addition dividuals, sounds clarkii emitted subordates by domcould 92

4 hen Mok: Sound Production nemonefishes Fig. 4. Spectrograms chirp sound Fig. 3., Fig. 3;, Fig. 3;, Fig. 3. Fig. 5. Oscillograms sgle pop sounds produced by mphiprion clarkii () frenatus ().

5 Japan. J. Ichthyol.35(1),1988 Fig. 6. Occurrence distribution pulse numbers pop sounds mphiprion clarkii frenatus. Fig. 7. Sonagrams chirp sounds preceeded by pop sound (dicated by an arrow) mphiprion clarkii () frenatu.s. ().

6 hen Molc: Sound Production nemonefishes Fig. 8. Oscillograms chirp sounds Fig. 3., Fig. 3;, Fig. 3;, Fig. 3. Fig. 9. Occurrence distribution pulse numbers chirp sound mphiprion clarkii (). frenatus (). N, number chirp sound analyzed.

7 魚類学 雑誌 Japan. J. Ichthyol.35(1),1988 D E Fig. 10. Tooth patterns cerabranchial plate mphiprion clarkii (), Irenatus (), perideraion (), oceliaris (D), Premnus biaculeatus (E). D E Fig. 11. Tooth patterns pharyngobranchial oth plates mphiprion clarkii (), frenatus (), perideraion (), ocellaris (D), Premnus biaculemus (E). ph2, oth plate second pharyngobranchial; ph3+4, oth plate third fourth pharyngobranchials. Fig. 12. Shakg sound emitted by an mphiprion 96 clarkii subordate. rrows pot this sound.

8 hen Mok: Sound Production nemonefishes with aggression, this difference may dicate species-specific variation aggression wards conspecifics. Difference has also been reported (e.g., Mariscal, 1972) on species-specific aggression shown durg terspecific teraction. thorough understg controllg facrs terspecific traspecific aggressions (e.g., terspecific traspecific socialities, predarprey selective pressure, capability on terspecific competition, utilization symbiotic host) relationship between se aggressive levels may lead a better knowledge anemonefish vocalization. cknowledgments We want thank Drs. Gerald R. llen Western ustralian Museum Richard N. Mariscal Florida State University for providg us with many important references. Thanks are also due anonymous reviewer for his or her helpful comments on manuscript. This paper is a portion master sis senior author submitted National Sun Yat-sen University, Republic ha for partial fulfillment master degree. Literature cited llen, G.R The anemonefishes. Their classification biology. T.F.H. Publ., Neptune ity, 288pp. hen, K The phylogenetic relationship four anemonefishes, mphiprion clarkii, frenatus, ocellaris, perideraion (Pisces, Pomacentridae) from ethological, morphological, biochemical approaches. Master Thesis, National Sun Yat-sen University, Kaohsiung, 112pp. Eibl-Eibesfeldt, I eobachtungen und Versuche an nemonefischen (mphiprion) der Malediven und der Nicobaren. Z. Tierpsychol., 17(1): Mariscal, R.N ehavior symbiotic fishes sea anemones. Pages H.E. Wn.L. Olla, eds. ehavior mare animals. Vol.2. Plenum Press, New York. Myrberg,, Jr. R.J. Riggio coustically mediated dividual recognition by a coral reef fish (Pomacentrus partitus). nim. ehav., 33: Myrberg,, Jr. J.Y. Shires Hearg damselfishes: an analysis signal detection among closely related species. J. omp. Physiol., 140: Myrberg,,Jr.,E. Spanier S.J.Ha Temporal patterng acoustical communication. Pages E.S. Reese F.J. Lighter, eds. ontrast behavior: adaptations aquatic terrestrial environment. John Wiley & Sons, New York. Schneider, H ioakustische Untersuchungen an nemonenfischen der Gattung mphiprion (Pisces). Z. Morph. Okol. Tiere., 53: Spanier, E spects species recognition by sound four species damselfishes, genus Eupomacentrus (Pisces: Pomacentridae). Z. Tierpsychol., 51: Verwey, J oral reef studies. I. The symbiosis between damselfishes sea anemones atavia ay. Treubia, 12(3-4): (Institute Mare iology, National Sun Yat-sen University, Kaohsiung, Taiwan 80424, Republic ha)