The Male Song of the Javan Silvery Gibbon (Hylobates Moloch)
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Contributions to Zoology, 74 (1/2) 1-25 (2005) The male song of the Javan silvery gibbon (Hylobates moloch) Thomas Geissmann1, Sylke Bohlen-Eyring2 and Arite Heuck2 1 Anthropological Institute, University Zürich-Irchel, Winterthurerstr. 190, CH-8057 Zürich, Switzerland; 2 Institute of Zoology, Tierärztliche Hochschule Hannover, Germany Keywords: Hylobates moloch, silvery gibbon, male song, individuality, calls, honest signal Abstract Contents This is the first study on the male song of the Javan silvery Introduction ....................................................................1 gibbon (Hylobates moloch), and the first quantitative Material and methods.....................................................3 evaluation of the syntax of male solo singing in any gibbon Study animals .............................................................3 species carried out on a representative sample of individuals. Recording and analysis equipment ...........................3 Because male gibbon songs generally exhibit a higher degree Acoustic terms and definitions ..................................3 of structural variability than female songs, the syntactical rules and the degree of variability in male singing have rarely Data collection............................................................3 been examined. In contrast to most other gibbon species, Statistics ......................................................................5 mated silvery gibbons do not appear to produce duet song Results.............................................................................5 bouts but solo song bouts only, and male singing is exceptio- Repertoire of note classes and types .........................5 nally rare, making this study particularly challenging. For the Frequency of note types.............................................8 present study, we tape-recorded and analysed several solo Inter-phrase interval .................................................12 song bouts of eight silvery gibbon males, including both wild Song motivation .......................................................12 and captive individuals. Based on their frequency characte- Phrase composition ..................................................13 ristics, song notes were classified into a total of 14 note Discussion.....................................................................17 types. These can be grouped into five groups (labelled A Male note repertoire .................................................17 through E). The proportions of the various note types were determined individually for successive 50-note sections Composition of the song bout .................................17 throughout the whole song bout. Based on changes in the Phrase syntax ............................................................17 proportion of different note classes and note types, we Individuality .............................................................18 roughly identify two phases in the male song: an introductory Comparison with male singing in other gibbons....18 phase, during which A and B1 notes are dominant, B3 notes Possible functions of male singing..........................19 are rare and C notes are absent, and a main song phase, New functional hypotheses......................................21 during which B3 or C notes are dominant. The occurrence Conclusions ..................................................................22 and the proportion of various types apparently differ among Acknowledgements......................................................22 individual males, however, both in the introductory and in References ....................................................................22 the main phase. We estimated song motivation by deter- Appendix ......................................................................24 mining the “number of notes per phrase” for each 50-note segment of the song bout. In each song bout, song motivation quickly increases during the introductory phase. Song motivation may exhibit strong fluctuations during the main Introduction phase of the song, but usually remains above values of 2 notes per phrase and thus above the values observed during the introductory phase. Males appear to exhibit individual The gibbons or small apes are distributed through- preferences in the order of different note types used in their out the tropical rain forests of South-east Asia phrases. Phrase structure was found to exhibit unusually low (Chivers, 2001; Geissmann, 1995; Marshall and degrees of stereotypy and high degrees of variability. As a Sugardjito, 1986). They live in monogamous, surprising finding of our study, male phrase variability both territorial family groups typically consisting of an within and between individuals appears to be higher in H. moloch than in most, perhaps all, other gibbon species. This adult pair and 1-3 immature offspring (Chivers, high variability appears to be a derived characteristic among 1977, 2001; Leighton, 1987). All species of the Hylobatidae. We discuss the implications of this finding gibbons are known to produce elaborate, loud, for the interpretation of song function and present new and long and stereotyped patterns of vocalisation testable functional hypotheses. Our study demonstrates that song function cannot be identified for “the gibbon”. Gibbon usually referred to as “songs” (Geissmann, 1993, songs appear to be multifunctional, and the relevance of 1995, 2000; Haimoff, 1984; Marshall and these functions appears to exhibit strong differences among Marshall, 1976). Preferentially, song bouts are gibbon species. produced in the early morning and have a duration of about 10-30 minutes. In most gibbon species, 2 T. Geissmann et al. – The Male Song of the Javan Silvery Gibbon mated pairs utter their songs in the form of well- male silvery gibbon song bouts occurring about coordinated duets. In addition, mated males of the once each week on average and usually before lar group (genus Hylobates) may also produce dawn. Male songs usually ended before the male solo songs (Geissmann, 2002a). females started their song bouts at around dawn. The Javan silvery gibbon (Hylobates moloch) is Acoustic individuality has been demonstrated one of only two gibbon species not known to to exist in several gibbon species (Dallmann and produce duet songs (the other non-duetting Geissmann, 2001a, b; Haimoff and Gittins, 1985; species being H. klossii) (Geissmann, 2002a). In Haimoff and Tilson, 1985), but these studies have addition, H. moloch is particularly unusual among been restricted to female singing. Female songs, hylobatids because most of the singing is as a rule, are more stereotyped than male songs produced by females (Geissmann and Nijman, (Geissmann, 1995, 2002a) and, therefore, easier to 2001; Kappeler, 1984a), and mated females analyse. appear to be the vocal “ ’representative’ of the The elusive male song of H. moloch has not family” (Kappeler, 1984a). In all other gibbons, previously been analysed quantitatively. All we either the opposite is true or the amounts of male know of the song is based on short verbal and female singing are about equal (Geissmann, descriptions supplemented with few short 2002a). sonagrams (Kappeler, 1984a; Geissmann and Because male singing is exceptionally rare in Nijman, 2001). An additional song of a captive silvery gibbons, studying these songs was male has been described by Haimoff (1983), but particularly challenging. During 130 days in this individual was later identified as a Bornean Western Java, Kappeler heard none of his five gibbon (H. muelleri) (Geissmann, 2002a). study group males produce a song and concluded In this study, we describe the syntax of the “it thus appears that territorial male moloch silvery gibbon male song based on tape-recorded gibbons do not sing” (Kappeler, 1984a, p. 386). songs of several captive and wild individuals. We The only male song Kappeler heard in his study compare the variability of male song structure area was produced by a solitary male singing on within and between individuals. Our results can the common territory boundary of two resident be used as a baseline for future studies on groups. Another similar song bout was heard individuality in male gibbon singing, because the outside of the study area. More recent obser- syntax and variability of male solo songs have vations in Central Java revealed that territorial apparently not been studied quantitatively in a males do sing, albeit rarely (Geissmann and representative sample of individuals for any other Nijman, 2001). These authors heard choruses of gibbon species. Table 1. Origin, number of individuals and song bouts, and recording information of the material analysed in this study. Recording a Recording Date Individual Author of Recording Duration of Completeness of Recording Song Bout Halimun 23 Nov. 1998 “Halimun” B. Merker 14 min 43 s No Howletts 1 17 Oct. 1988 “Omar” T. Geissmann 38 min 46 s Yes Howletts 2 18 June 1995 “Hilo” T. Geissmann 35 min 52 s Yes b Howletts 3 28 Aug. 1999 “Lupau” S. Bohlen-Eyring 14 min 42 s Yes Jakarta 1a 8 June 1978 “Jakarta 1” M. Kappeler 7 min 31 s No Jakarta 1b 7 June 1978 “Jakarta 1” M. Kappeler 8 min 26 s No Jakarta 2 28 Jan. 2001 “Jakarta 2” T. Geissmann 40 min 45 s No Kalejetan 25 Aug.1978 “Kalejetan” M. Kappeler 2 min 17 s No Munich 1a 21 June 1988 “Paul” T. Geissmann 12 min 15 s Yes c Munich 1b 17 Oct. 1991 “Paul” T. Geissmann 21 min 03 s Yes Munich 1c 28 April 1995 “Paul” T. Geissmann 20 min 02 s Yes a Recording