Formants provide honest acoustic cues to body size in American alligators
Open accessAbstract
In many vertebrates, acoustic cues to body size are encoded in resonance frequencies of the vocal tract (“formants”), rather than in the rate of tissue vibration in the sound source (“pitch”). Anatomical constraints on the vocal tract’s size render formants honest cues to size in many bird and mammal species, but it is not clear whether this correlation evolved convergently in these two clades, or whether it is widespread among amniotes (mammals, birds, and non-avian reptiles). We investigated the potential for honest acoustic cues in the bellows of adult American alligators and found that formant spacing provided highly reliable cues to body size, while presumed correlates of the source signal did not. These findings held true for both sexes and for all bellows whether produced in or out of water. Because birds and crocodilians are the last extant Archosaurians and share common ancestry with all extinct dinosaurs, our findings support the hypothesis that dinosaurs used formants as cues to body size. The description of formants as honest signals in a non-avian reptile combined with previous evidence from birds and mammals strongly suggests that the principle of honest signalling via vocal tract resonances may be a broadly shared trait among amniotes.
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Vocalisations such as growls, roars, and bellows are produced by air pressure travelling through the laryngeal vocal folds causing them to vibrate through the trachea, with resonance produced by the vocal tract.
“These include vocalisations such as growls, roars (in crocodiles) and bellows (in alligators), all of which are produced by air pressure travelling through the laryngeal vocal folds causing them to vibrate through the trachea, with resonance produced by the vocal tract (Riede et al. 2011; Reber et al. 2017).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
The Introduction identifies bellowing and roaring as crocodilian vocalisations, and the Discussion supports vocal-tract resonance in alligator bellows. But the Discussion explicitly says it remains unclear whether the vibrating tissue is the vocal folds, so the cited paper does not support that part of the proposed mechanism. The supplied passages have no page numbers. (LLM, conf=0.88)
Experimental — accuracy not yet measuredThis size-frequency relationship is based on whether vocal fold size (the main determinant of frequency) correlates with overall body size, which may not be the case for all species.
“This size‐frequency relationship is based on whether vocal fold size (the main determinant of frequency) correlates with overall body size, which may not be the case for all species (Reber et al. 2017; Jensen et al. 2024).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
The Introduction (page unavailable) says that fundamental frequency reflects the rate of source-tissue vibration and that, in many species, it has no apparent link to body size, possibly because vibrating-tissue size is not tightly constrained by anatomy. This supports the claim that a size–frequency relationship need not hold across species, but the paper does not establish that vocal-fold size is the main determinant of frequency. (LLM, conf=0.81)
Experimental — accuracy not yet measuredSex appears to significantly predict some of the acoustic parameters of different call types, and dominant frequency (along with all formant-related parameters) in American alligator bellows is higher in females than in males, with this trend consistent across studies.
“Sex appears to significantly predict some of the acoustic parameters of different call types.”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
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Experimental — accuracy not yet measuredVocalisations are composed of a fundamental frequency, harmonics, and for some species formants, all of which potentially modulate in frequency and amplitude across time.
“Vocalisations, for example, are composed of a fundamental frequency, harmonics (Vergne et al. 2012), and for some species formants (Reber et al. 2015, 2017), all of which potentially modulate in frequency and amplitude across time (Vergne et al. 2012).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
Formants exist in the bellows produced by the Chinese alligator and likely also the American alligator.
“Such formants exist in the bellows produced by the Chinese alligator (Alligator sinensis, Fauvel 1879) (Reber et al. 2015) and likely also the American alligator (Alligator mississippiensis, Daudin 1801) (Reber et al. 2017).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
Formants are produced from the resonance of the vocal tract and therefore scale with body size, and have been shown to contain individual identity, predict sex in American alligator bellows, and likely contain information pertaining to body size and fitness in this species.
“Formants are produced from the resonance of the vocal tract and therefore scale with body size (Reber et al. 2017).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Reber et al. 2017
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Cites
Social Signals and Behaviors of Adult Alligators and Crocodiles
GARRICK, L. D., & LANG, J. W. (1977) · American Zoologist · doi:10.1093/icb/17.1.225 · Cited by 193 · see citing works
Signalling among relatives. I. Is costly signalling <i>too</i> costly?
Bergstrom, C. T., & Lachmann, M. (1997) · Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rstb.1997.0041 · Cited by 89 · see citing works
Size does matter: crocodile mothers react more to the voice of smaller offspring
Chabert, T., Colin, A., Aubin, T., Shacks, V., Bourquin, S. L., Elsey, R. M., Acosta, J. G., & Mathevon, N. (2015) · Scientific Reports · doi:10.1038/srep15547 · Cited by 60 · see citing works
The peacock train does not handicap cursorial locomotor performance
Thavarajah, N. K., Tickle, P. G., Nudds, R. L., & Codd, J. R. (2016) · Scientific Reports · doi:10.1038/srep36512 · Cited by 25 · see citing works
Flagged first, then by citations
Research works printed but not linked: 54
- [2]Clutton-Brock TH Albon SD Roaring of red deer and the evolution of honest advertisement Behaviour 1979 69 145 170 10.1163/156853979X00449 · DOI
- [3]Charlton BD Vocal cues to male androgen levels in giant pandas Biol. Lett. 2010 7 71 74 10.1098/rsbl.2010.0582 20810426 PMC3030892 · DOI
- [4]Harris TR Fitch WT Goldstein LM Fashing PJ Black and White colobus monkey (Colobus guereza) roars as a source of both honest and exaggerated information about body mass Ethology 2006 112 911 920 10.1111/j.1439-0310.2006.01247.x · DOI
- [5]Smith JM Price GR The logic of animal conflict Nature 1973 246 15 18 10.1038/246015a0 · DOI
- [7]Zahavi A Mate selection—A selection for a handicap J. Theor. Biol. 1975 53 205 214 10.1016/0022-5193(75)90111-3 1195756 · DOI · Paper page
- [8]Grafen A Biological signals as handicaps J. Theor. Biol. 1990 144 517 5468 10.1016/S0022-5193(05)80088-8 2402153 · DOI · Paper page
- [9]Siller S A note on errors in Grafen’s strategic handicap models J. Theor. Biol. 1998 195 413 417 10.1006/jtbi.1998.0693 9826494 · DOI
- [10]Kotiaho JS Costs of sexual traits: a mismatch between theoretical considerations and empirical evidence Biol. Rev. 2001 76 365 376 10.1017/S1464793101005711 11569789 · DOI
- [11]Számadó S The cost of honesty and the fallacy of the handicap principle Anim. Behav. 2011 81 3 10 10.1016/j.anbehav.2010.08.022 · DOI
- [13]Silk JB Kaldor E Boyd R Cheap talk when interests conflict Animal Behaviour 2000 59 423 432 10.1006/anbe.1999.1312 10675265 · DOI
- [15]Bergstrom CT Lachmann M Signaling among relatives. III Talk is cheap. PNAS 1998 95 5100 5105 9560235 PMC20220
- [16]Maynard-Smith, J. & Harper, D. Animal signals. (Oxford University Press, 2003).
- [17]Fitch, W. T. & Hauser, M. D. In Acoustic Communication Vol. 16 (eds Simmons, A. M., Popper, A. N. & Fay, R. R.) 65-137 (Springer, 2002).
- [18]Riechert SE Games spiders play: Behavioral variability in territorial disputes Behav. Ecol. Sociobiol. 1978 3 135 162 10.1007/BF00294986 · DOI
- [19]Reby D McComb K Anatomical constraints generate honesty: acoustic cues to age and weight in the roars of red deer stags Anim. Behav. 2003 65 519 530 10.1006/anbe.2003.2078 · DOI
- [20]Fitch WT The phonetic potential of nonhuman vocal tracts: Comparative cineradiographic observations of vocalizing animals Phonetica 2000 57 205 218 10.1159/000028474 10992141 · DOI
- [21]Fitch WT Vocal tract length and formant frequency dispersion correlate with body size in rhesus macaques J. Acoust. Soc. Am. 1997 102 1213 1222 10.1121/1.421048 9265764 · DOI
- [23]Taylor AM Reby D The contribution of source-filter theory to mammal vocal communication research J. Zool. 2010 280 221 236 10.1111/j.1469-7998.2009.00661.x · DOI · Paper page
- [24]Titze, I. R. Principles of voice production. (Prentice Hall, 1994).
- [25]Morton ES On the occurrence and significance of motivation - Structural rules in some bird and mammal sounds Am. Nat. 1977 111 855 869 10.1086/283219 · DOI
- [26]Lass NJ Brown WS Correlational study of speakers’ heights, weights, body surface areas, and speaking fundamental frequencies J. Acoust. Soc. Am. 1978 63 1218 1220 10.1121/1.381808 649880 · DOI
- [27]Masataka N Lack of correlation between body size and frequency of vocalizations in young female Japanese macaques (Macaca fuscata) Folia Primatol. 1994 63 115 118 10.1159/000156803 7813976 · DOI
- [28]Rendall D Kollias S Ney C Lloyd P Pitch (F0) and formant profiles of human vowels and vowel-like baboon grunts: The role of vocalizer body size and voice-acoustic allometry J. Acoust. Soc. Am. 2005 117 944 955 10.1121/1.1848011 15759713 · DOI
- [29]Pfefferle D West PM Grinnell J Packer C Fischer J Do acoustic features of lion, Panthera leo, roars reflect sex and male condition? J. Acoust. Soc. Am. 2007 121 3947 3953 10.1121/1.2722507 17552741 · DOI
- [30]Fitch WT Hauser MD Vocal production in nonhuman primates: acoustics, physiology, and functional constraints on “honest” advertisement Am. J. Primatol. 1995 37 191 219 10.1002/ajp.1350370303 31936952 · DOI
- [31]Reby D Red deer stags use formants as assessment cues during intrasexual agonistic interactions Proc. R. Soc. B. 2005 272 941 947 10.1098/rspb.2004.2954 16024350 PMC1564087 · DOI
- [32]Charlton BD Reby D The evolution of acoustic size exaggeration in terrestrial mammals Nat. Commun. 2016 7 8 10.1038/ncomms12739 PMC5025854 27598835 · DOI
- [33]Fitch WT Acoustic exaggeration of size in birds via tracheal elongation: comparative and theoretical analyses J. Zool. 1999 248 31 48 10.1111/j.1469-7998.1999.tb01020.x · DOI
- [34]Hinds DS Calder WA Tracheal dead space in the respiration of birds Evolution 1971 25 429 440 10.2307/2406936 28563117 · DOI
- [35]Jones MR Witt CC Migrate small, sound big: functional constraints on body size promote tracheal elongation in cranes J. Evol. Biol. 2014 27 1256 1264 10.1111/jeb.12397 24800977 · DOI
- [36]Miller EH Williams J Jamieson SE Gilchrist HG Mallory ML Allometry, bilateral asymmetry and sexual differences in the vocal tract of common eiders Somateria mollissima and king eiders S. spectabilis J. Avian Biol. 2007 38 224 233 10.1111/j.0908-8857.2007.03695.x · DOI
- [38]Weissengruber GE Forstenpointner G Peters G Kubber-Heiss A Fitch WT Hyoid apparatus and pharynx in the lion (Panthera leo), jaguar (Panthera onca), tiger (Panthera tigris), cheetah (Acinonyx jubatus) and domestic cat (Felis silvestris f. catus) J. Anat. 2002 201 195 209 10.1046/j.1469-7580.2002.00088.x 12363272 PMC1570911 · DOI
- [39]Fitch WT Reby D The descended larynx is not uniquely human Proc. R. Soc. B. 2001 268 1669 1675 10.1098/rspb.2001.1704 11506679 PMC1088793 · DOI
- [40]Rand AS Dudley R Frogs in helium: The anuran vocal sac is not a cavity resonator Physiol. Zool. 1993 66 793 806 10.1086/physzool.66.5.30163824 · DOI
- [41]Gans C Maderson PFA Sound Producing Mechanisms in Recent Reptiles: Review and Comment Am. Zool. 1973 13 1195 1203 10.1093/icb/13.4.1195 · DOI
- [43]Vergne AL Pritz MB Mathevon N Acoustic communication in crocodilians: from behaviour to brain Biol. Rev. 2009 84 391 411 10.1111/j.1469-185X.2009.00079.x 19659884 · DOI · Paper page
- [44]Vergne AL Mathevon N Crocodile egg sounds signal hatching time Curr. Biol. 2008 18 513 514 10.1016/j.cub.2008.04.011 18579090 · DOI
- [45]Vergne AL Aubin T Taylor P Mathevon N Acoustic signals of baby black caimans Zoology 2011 114 313 320 10.1016/j.zool.2011.07.003 21978842 · DOI
- [47]Brazaitis P Watanabe ME Crocodilian behaviour: a window to dinosaur behaviour? Hist. Biol. 2011 23 73 90 10.1080/08912963.2011.560723 · DOI
- [48]Tucker AD McCallum HI Limpus CJ McDonald KR Sex-biased dispersal in a long-lived polygynous reptile (Crocodylus johnstoni) Behav. Ecol. Sociobiol. 1998 44 85 90 10.1007/s002650050519 · DOI
- [49]Webb GJW Manolis SC Crocodylus johnstoni in the McKinlay River, N.T. V. Abnormalities and injuries Aust. Wildlife Res. 1983 10 407 420 10.1071/WR9830407 · DOI
- [52]Vliet KA Social displays of the American alligator (Alligator mississippiensis) Am. Zool. 1989 29 1019 1031 10.1093/icb/29.3.1019 · DOI · Paper page
- [53]Reber SA Nishimura T Janisch J Robertson M Fitch WT A Chinese alligator in heliox: formant frequencies in a crocodilian J. Exp. Biol. 2015 218 2442 2447 10.1242/jeb.119552 26246611 PMC4528706 · DOI · Paper page
- [54]Brunell AM Delaney JP Spratt RG Carbonneau DA Waller JE Record total lengths of the American alligator in Florida Southeast. Nat. 2013 12 9 17 10.1656/058.012.0411 · DOI
- [55]Woodward AR White JH Linda SB Maximum size of the alligator (Alligator mississippiensis) J. Herpetol. 1995 29 507 513 10.2307/1564733 · DOI
- [56]Erickson GM Lappin AK Parker T Vliet KA Comparison of bite-force performance between long-term captive and wild American alligators (Alligator mississippiensis) J. Zool. 2004 262 21 28 10.1017/S0952836903004400 · DOI · Paper page
- [58]Leitch DB Catania KC Structure, innervation and response properties of integumentary sensory organs in crocodilians J. Exp. Biol. 2012 215 4217 4230 10.1242/jeb.076836 23136155 PMC4074209 · DOI
- [59]Fitch WT Kelley JP Perception of vocal tract resonances by whooping cranes Grus americana Ethology 2000 106 559 574 10.1046/j.1439-0310.2000.00572.x · DOI
- [60]Charlton BD Ellis WAH Larkin R Fitch WT Perception of size-related formant information in male koalas (Phascolarctos cinereus) Anim. Cogn. 2012 15 999 1006 10.1007/s10071-012-0527-5 22740017 · DOI
- [61]Charlton BD Reby D McComb K Female perception of size-related formant shifts in red deer Cervus elaphus. Anim. Behav. 2007 74 707 714 10.1016/j.anbehav.2006.09.021 · DOI
- [62]Hedges SB Poling LL A Molecular Phylogeny of Reptiles Science 1999 283 998 1001 10.1126/science.283.5404.998 9974396 · DOI
- [63]Weishampel DB Acoustic analyses of potential vocalization in Lambeosaurine dinosaurs (Reptilia, Ornithischia) Paleobiology 1981 7 252 261 10.1017/S0094837300004036 · DOI
- [65]Halekoh U Hojsgaard S Kenward-Roger approximation and parametric bootstrap methods for tests in linear mixed models - The R package pbkrtest J. Stat. Softw. 2014 59 1 32 10.18637/jss.v059.i09 · DOI · Paper page
- [66]Mundry R Fischer J Use of statistical programs for nonparametric tests of small samples often leads to incorrect P values: examples from Animal Behaviour Anim. Behav. 1998 56 256 259 10.1006/anbe.1998.0756 9710485 · DOI
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- Authors
- Reber, S. A., Janisch, J., Torregrosa, K., Darlington, J., Vliet, K. A., & Fitch, W. T.
- Affiliations
- University of Vienna (AT)
- Wildlife Conservation Society (US)
- University of Florida (US)
- Corresponding author
- Reber, Stephan A.
- Fitch, W. Tecumseh
- Venue
- Scientific Reports
- Published
- 2017-05-12
- Volume
- 7
- Issue
- 1
- Pages
- 1816-1816
- ISSN
- 2045-2322 (ISSN-L)
- PMC
- PMC5431764
- OpenAlex
- W2612365460
- Cited by
- 40
- References
- 4
- Citations by year
- 2026: 2 · 2025: 7 · 2024: 3 · 2023: 4 · 2022: 4 · 2021: 7 · 2020: 7 · 2019: 2 · 2018: 4
- FWCI
- 3.30
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Scientific Reports (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Phaidra (Universität Wien) (submittedVersion, cc-by-nc-nd)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Lund University Publications (Lund University) (submittedVersion)
- Funders
- Austrian Science Fund (W-1234-G17, W 1234, 230604)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Physical Sciences › Environmental Science › Ecology › Marine animal studies overview
- Keywords
- formants, body size, honest signaling, Alligator mississippiensis, crocodilians, vocal tract resonance, bellows, acoustic communication, amniotes, dinosaurs, convergent evolution, sex differences
- MeSH terms
- Alligators and Crocodiles, Animals, Cues, Female, Male, Vocalization, Animal, Body Size
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