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Complex signal function: developing a framework of testable hypotheses

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Abstract

The basic building blocks of communication are signals, assembled in various sequences and combinations, and used in virtually all inter- and intra-specific interactions. While signal evolution has long been a focus of study, there has been a recent resurgence of interest and research in the complexity of animal displays. Much past research on signal evolution has focused on sensory specialists, or on single signals in isolation, but many animal displays involve complex signaling, or the combination of more than one signal or related component, often serially and overlapping, frequently across multiple sensory modalities. Here, we build a framework of functional hypotheses of complex signal evolution based on content-driven (ultimate) and efficacy-driven (proximate) selection pressures (sensu Guilford and Dawkins 1991). We point out key predictions for various hypotheses and discuss different approaches to uncovering complex signal function. We also differentiate a category of hypotheses based on inter-signal interactions. Throughout our review, we hope to make three points: (1) a complex signal is a functional unit upon which selection can act, (2) both content and efficacy-driven selection pressures must be considered when studying the evolution of complex signaling, and (3) individual signals or components do not necessarily contribute to complex signal function independently, but may interact in a functional way.

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References

  • Acquistapace P, Aquiloni L, Hazlett BA, Gherardi F (2002) Multimodal communication in crayfish: sex recognition during mate search by male Austropotamobius pallipes. Can J Zool 80:2041–2045

    Article  Google Scholar 

  • Andersson M (1982) Female choice selects for extreme tail length in a Widowbird. Nature 299:818–820

    Google Scholar 

  • Badyaev AV, Hill GE (2000) Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration. Biol J Linn Soc 69:153–172

    Article  Google Scholar 

  • Bailey WJ (1991) Acoustic behaviour of insects: an evolutionary perspective. Chapman and Hall, London

    Google Scholar 

  • Bailey WJ, Cunningham RJ, Lebel L (1990) Song power, spectral distribution and female phonotaxis in the bushcricket Requena-Verticalis (Tettigoniidae, Orthoptera) - active female choice or passive attraction. Anim Behav 40:33–42

    Google Scholar 

  • Balakrishnan R, Helversen D von, Helversen O von (2001) Song pattern recognition in the grasshopper Chorthippus biguttulus: the mechanism of syllable onset and offset detection. J Comp Physiol A 187:255–264

    Article  CAS  PubMed  Google Scholar 

  • Basolo AL (1990) Female preference predates the evolution of the sword in swordtail fish. Science 250:808–810

    Google Scholar 

  • Baurecht D, Barth FG (1992) Vibratory communication in spiders .1. Representation of male courtship signals by female vibration receptor. J Comp Physiol A 171:231–243

    Article  Google Scholar 

  • Birkhead TR, Fletcher F, Pellatt EJ (1998) Sexual selection in the zebra finch Taeniopygia guttata: condition, sex traits and immune capacity. Behav Ecol Sociobiol 44:179–191

    Article  Google Scholar 

  • Bischoff RJ, Gould JL, Rubenstein DI (1985) Tail size and female choice in the guppy (Poecilia reticulata). Behav Ecol Sociobiol 17:253–255

    Article  Google Scholar 

  • Blanco G, De la Puente J (2002) Multiple elements of the black-billed magpie’s tail correlate with variable honest information on quality in different age/sex classes. Anim Behav 63:217–225

    Article  Google Scholar 

  • Borgia G, Presgraves DC (1998) Coevolution of elaborated male display traits in the spotted bowerbird: an experimental test of the threat reduction hypothesis. Anim Behav 56:1121–1128

    Article  PubMed  Google Scholar 

  • Bradbury JW, Vehrencamp SL (1998) Principles of animal communication. Sinauer, Sunderland, Mass.

  • Brodie ED (1992) Correlational selection for color pattern and antipredator behavior in the garter snake Thamnophis ordinoides. Evolution 46:1284–1298

    Google Scholar 

  • Brooks R, Caithness N (1995) Does a males attractiveness to a female depend on her previous experience. S Afr J Sci 91:156–158

    Google Scholar 

  • Buchholz R (1991) Older males have bigger knobs—Correlates of ornamentation in 2 species of curassow. Auk 108:153–160

    Google Scholar 

  • Candolin U (2003) The use of multiple cues in mate choice. Biol Rev 78:575–595

    Article  PubMed  Google Scholar 

  • Candolin U, Reynolds JD (2001) Sexual signaling in the European bitterling: females learn the truth by direct inspection of the resource. Behav Ecol 12:407–411

    Article  Google Scholar 

  • Coleman SW, Patricelli GL, Borgia G (2004) Variable female preferences drive complex male displays. Nature 428:742–745

    Article  CAS  PubMed  Google Scholar 

  • Cooper BG, Goller F (2004) Multimodal signals: enhancement and constraint of song motor patterns by visual display. Science 303:544–546

    Article  CAS  PubMed  Google Scholar 

  • Cott HB (1940) Adaptive coloration in animals. Methuen, London

  • Cynx J, Lewis R, Tavel B, Tse H (1998) Amplitude regulation of vocalizations in noise by a songbird, Taeniopygia guttata. Anim Behav 56:107–113

    Article  PubMed  Google Scholar 

  • De caprona MDC, Ryan MJ (1990) Conspecific mate recognition in swordtails, Xiphophorus nigrensis and X. pygmaeus (Poeciliidae)—Olfactory and visual cues. Anim Behav 39:290–296

    Google Scholar 

  • Doucet SM, Montgomerie R (2003) Multiple sexual ornaments in satin bowerbirds: ultraviolet plumage and bowers signal different aspects of quality. Behav Ecol 14:503–509

    Article  Google Scholar 

  • Draganoiu TI, Nagle L, Kreutzer M (2002) Directional female preference for an exaggerated male trait in canary (Serinus canaria) song. Proc R Soc Lond B Biol Sci 269:2525–2531

    Article  PubMed  Google Scholar 

  • Dukas R (2002) Behavioural and ecological consequences of limited attention. Philos Trans R Soc Lond B Biol Sci 357:1539–1547

    Article  PubMed  Google Scholar 

  • Dyer FC (2002) The biology of the dance language. Annu Rev Entomol 47:917–949

    Article  CAS  PubMed  Google Scholar 

  • Edmunds M (1974) Defense in animals. Longman, Harlow

  • Endler JA (1992) Signals, signal conditions, and the direction of evolution. Am Nat 139:S125-S153

    Article  Google Scholar 

  • Endler JA, Thery M (1996) Interacting effects of lek placement, display behavior, ambient light, and color patterns in three neotropical forest-dwelling birds. Am Nat 148:421–452

    Article  Google Scholar 

  • Endler JA, Basolo AL, Glowacki S, Zerr J (2001) Variation in response to artificial selection for light sensitivity in guppies (Poecilia reticulata). Am Nat 158:36–48

    Article  Google Scholar 

  • Ewing A (1989) Arthropod bioacoustics: neurobiology and behavior. Cornell University, Ithaca

    Google Scholar 

  • Fleishman LJ (2000) Signal function, signal efficiency and the evolution of anoline lizard dewlap color. In: Epsmark Y, Amundsen T GR (eds) Animal Signals: signalling and signal design in animal communication. Tapir Academic, Trondheim, pp 209–236

  • Fox D (1976) Animal biochromes and structural colors. University of California Press, Berkeley, Calif.

  • Gerhardt HC (1991) Female mate choice in treefrogs—static and dynamic acoustic criteria. Anim Behav 42:615–635

    Google Scholar 

  • Gerhardt HC, Huber F (2002) Acoustic communication in insects and anurans. University of Chicago Press, Chicago

  • Glassey B, Forbes S (2002) Muting individual nestlings reduces parental foraging for the brood. Anim Behav 63:779–786

    Article  Google Scholar 

  • Gong A, Gibson RM (1996) Reversal of a female preference after visual exposure to a predator in the guppy, Poecilia reticulata. Anim Behav 52:1007–1015

    Article  Google Scholar 

  • Grahn M, Vonschantz T (1994) Fashion and age in pheasants—age-differences in mate choice. Proc R Soc Lond B Biol Sci 255:237–241

    Google Scholar 

  • Grether GF (1996) Sexual selection and survival selection on wing coloration and body size in the rubyspot damselfly Hetaerina americana. Evolution 50:1939–1948

    Google Scholar 

  • Grether GF, Hudon J, Millie DF (1999) Carotenoid limitation of sexual coloration along an environmental gradient in guppies. Proc R Soc Lond B Biol Sci 266:1317–1322

    Article  CAS  Google Scholar 

  • Guilford T, Dawkins MS (1991) Receiver psychology and the evolution of animal signals. Anim Behav 42:1–14

    Google Scholar 

  • Halliday TR, Slater PJB (1983) Animal behaviour, vol 2. Freeman, New York

  • Hankison SJ, Morris MR (2002) Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences. Behav Ecol Sociobiol 51:140–145

    Article  Google Scholar 

  • Hasson O (1989) Amplifiers and the handicap principle in sexual selection: a different emphasis. Proc R Soc Lond B Biol Sci 235

    Google Scholar 

  • Hasson O (1991) Sexual displays as amplifiers—practical examples with an emphasis on feather decorations. Behav Ecol 2:189–197

    Google Scholar 

  • Hasson O (1994) Cheating signals. J Theor Biol 167:223–238

    Article  Google Scholar 

  • Hasson O (1997) Towards a general theory of biological signaling. J Theor Biol 185:139–156

    Article  PubMed  Google Scholar 

  • Hauser M (1996) The evolution of communication. MIT Press, Cambridge, Mass.

  • Hebets EA (2002) The evolution and function of complex signaling in spider courtship behavior. Ecology and evolutionary biology. PhD dissertation, University of Arizona, Tucson, Ariz.

  • Hebets E (2003) Subadult experience influences adult mate choice in an arthropod: exposed female wolf spiders prefer males of a familiar phenotype. Proc Natl Acad Sci USA 100:13390–13395

    Article  CAS  PubMed  Google Scholar 

  • Hebets EA (2004) Attention-altering interaction in the multimodal courtship display of the wolf spider Schizocosa uetzi. Behav Ecol (in press)

  • Hebets EA, Uetz GW (1999) Female responses to isolated signals from multimodal male courtship displays in the wolf spider genus Schizocosa (Araneae : Lycosidae). Anim Behav 57:865–872

    Article  PubMed  Google Scholar 

  • Hebets EA, Uetz GW (2000) Leg ornamentation and the efficacy of courtship display in four species of wolf spider (Araneae : Lycosidae). Behav Ecol Sociobiol 47:280–286

    Article  Google Scholar 

  • Hebets EA, Stratton GE, Miller GL (1996) Habitat and courtship behavior of the wolf spider Schizocosa retrorsa (Banks) (Araneae, Lycosidae). J Arachnol 24:141–147

    Google Scholar 

  • Holland B, Rice WR (1998) Perspective: chase-away sexual selection: antagonistic seduction versus resistance. Evolution 52:1–7

    Google Scholar 

  • Hölldobler B (1999) Multimodal signals in ant communication. J Comp Physiol A 184:129–141

    Article  Google Scholar 

  • Hughes M (1996) The function of concurrent signals: visual and chemical communication in snapping shrimp. Anim Behav 52:247–257

    Article  Google Scholar 

  • Jackson J, Ingram WI, Campbell HW (1976) The dorsal pigmentation pattern of snakes as an antipredator strategy: a multivariate approach. Am Nat 110:1029–1053

    Article  Google Scholar 

  • Jacobs GH, Neitz J, Neitz M (1993) Genetic basis of polymorphism in the color vision of platyrrhine monkeys. Vision Res 33:269–274

    Article  CAS  PubMed  Google Scholar 

  • Jawor JM, Gray N, Beall SM, Breitwisch R (2004) Multiple ornaments correlate with aspects of condition and behaviour in female northern cardinals, Cardinalis cardinalis. Anim Behav 67:875–882

    Article  Google Scholar 

  • Jetz W, Rowe C, Guilford T (2001) Non-warning odors trigger innate color aversions—as long as they are novel. Behav Ecol 12:134–139

    Article  Google Scholar 

  • Johnstone RA (1996) Multiple displays in animal communication: ‘backup signals’ and ‘multiple messages’. Philos Trans R Soc Lond B Biol Sci 351:329–338

    Google Scholar 

  • Kaye H, Mackintosh NJ, Rothschild M (1989) Odour of pyrazine potentiates an association between environmental cues and unpalatable taste. Anim Behav 37:563–568

    Google Scholar 

  • Leal M, Fleishman LJ (2004) Differences in visual signal design and detectability between allopatric populations of Anolis lizards. Am Nat 163:26–39

    Article  PubMed  Google Scholar 

  • Lengagne T, Slater PJB (2002) The effects of rain on acoustic communication: tawny owls have good reason for calling less in wet weather. Proc R Soc Lond B Biol Sci 269:2121–2125

    Article  PubMed  Google Scholar 

  • Lengagne T, Aubin T, Lauga J, Jouventin P (1999) How do king penguins (Aptenodytes patagonicus) apply the mathematical theory of information to communicate in windy conditions? Proc R Soc Lond B Biol Sci 266:1623–1628

    Article  Google Scholar 

  • Ligon JD, Zwartjes PW (1995) Ornate plumage of male red junglefowl does not influence mate choice by females. Anim Behav 49:117–125

    Article  Google Scholar 

  • Lindstrom L, Rowe C, Guilford T (2001) Pyrazine odour makes visually conspicuous prey aversive. Proc R Soc Lond B Biol Sci 268:159–162

    Article  CAS  PubMed  Google Scholar 

  • Lloyd JE (1971) Bioluminescent communication in insects. Annu Rev Entomol 16:97–122

    Article  Google Scholar 

  • Macaluso E, Frith CD, Driver J (2000) Modulation of human visual cortex by crossmodal spatial attention. Science 289:1206–1208

    Article  CAS  PubMed  Google Scholar 

  • Marchetti K (1998) The evolution of multiple male traits in the yellow-browed leaf warbler. Anim Behav 55:361–376

    Article  PubMed  Google Scholar 

  • Markl H (1983) Vibrational communication. In: Huber R, Markl H (eds) Neuroethology and behavioral physiology. Springer, Berlin Heidelberg New York, pp 332–353

  • Marples NM, Roper TJ (1996) Effects of novel colour and smell on the response of naive chicks towards food and water. Anim Behav 51:1417–1424

    Article  Google Scholar 

  • Marples NM, Van Veelen W, Brakefield PM (1994) The relative importance of colour, taste and smell in the protection of an aposematic insect Coccinella septempunctata. Anim Behav 48:967–974

    Article  Google Scholar 

  • Maynard Smith J, Harper DGC (1995) Animal signals: models and terminology. J Theor Biol 177:305–311

    Article  Google Scholar 

  • McClintock WJ, Uetz GW (1996) Female choice and pre-existing bias: visual cues during courtship in two Schizocosa wolf spiders (Araneae: Lycosidae). Anim Behav 52:167–181

    Article  Google Scholar 

  • McDonald CG, Hawryshyn CW (1995) Intraspecific variation of spectral sensitivity in threespine stickleback (Gasterosteus aculeatus) from different photic regimes. J Comp Physiol A 176:255–260

    Article  Google Scholar 

  • McGraw KJ, Hill GE (2000) Differential effects of endoparasitism on the expression of carotenoid- and melanin-based ornamental coloration. Proc R Soc Lond B Biol Sci 267:1525–1531

    Article  CAS  PubMed  Google Scholar 

  • McLennan DA (2003) The importance of olfactory signals in the gasterosteid mating system: sticklebacks go multimodal. Biol J Linn Soc 80:555–572

    Article  Google Scholar 

  • Meredith MA, Stein BE (1996) Spatial determinants of multisensory integration in cat superior colliculus neurons. J Neurophysiol 75:1843–1857

    CAS  PubMed  Google Scholar 

  • Møller AP, Pomiankowski A (1993) Why have birds got multiple sexual ornaments. Behav Ecol Sociobiol 32:167–176

    Google Scholar 

  • Møller AP, Thornhill R (1998) Bilateral symmetry and sexual selection: a meta-analysis. Am Nat 151:174–192

    Article  Google Scholar 

  • Narins PM, Capranica RR (1976) Sexual differences in auditory-system of tree frog Eleutherodactylus coqui. Science 192:378–380

    CAS  PubMed  Google Scholar 

  • Partan S, Marler P (1999) Behavior—communication goes multimodal. Science 283:1272–1273

    Article  CAS  PubMed  Google Scholar 

  • Patricelli GL, Uy JAC, Walsh G, Borgia G (2002) Male displays adjusted to female’s response—macho courtship by the satin bowerbird is tempered to avoid frightening the female. Nature 415:279–280

    Article  CAS  PubMed  Google Scholar 

  • Patricelli GL, Uy JAC, Borgia G (2003) Multiple male traits interact: attractive bower decorations facilitate attractive behavioural displays in satin bowerbirds. Proc R Soc Lond B Biol Sci 270:2389–2395

    Article  PubMed  Google Scholar 

  • Petrie M, Halliday T, Sanders C (1991) Peahens prefer peacocks with elaborate trains. Anim Behav 41:323–331

    Google Scholar 

  • Pfennig KS (1998) The evolution of mate choice and the potential for conflict between species and mate-quality recognition. Proc R Soc Lond B Biol Sci 265:1743–1748

    Article  Google Scholar 

  • Podos J (1997) A performance constraint on the evolution of trilled vocalizations in a songbird family (Passeriformes:Emberizidae). Evolution 51:537–551

    Google Scholar 

  • Reid ML, Weatherhead PJ (1990) Mate-choice criteria of Ipswich sparrows—the importance of variability. Anim Behav 40:538–544

    Google Scholar 

  • Ronacher B, Rudiger K (2000) Temporal integration vs. parallel processing: coping with the variability of neuronal messages in directional hearing of insects. Eur J Neurosci 12:2147–2156

    Article  CAS  PubMed  Google Scholar 

  • Ronacher B, Franz A, Wohlgemuth S, Hennig R (2004) Variability of spike trains and the processing of temporal patterns of acoustic signals—problems, constraints, and solutions. J Comp Physiol A 190:257–277

    Article  CAS  Google Scholar 

  • Roper TJ, Marples NM (1997) Odour and colour as cues for taste-avoidance learning in domestic chicks. Anim Behav 53:1241–1250

    Article  PubMed  Google Scholar 

  • Rosenthal GG, Rand AS, Ryan MJ (2004) The vocal sac as a visual cue in anuran communication: an experimental analysis using video playback. Anim Behav 68:55–58

    Article  Google Scholar 

  • Rothschild M (1961) Defensive odours and Mullerian mimicry among insects. Trans R Entomol Soc Lond 113:101–121

    Google Scholar 

  • Rothschild M, Haskell PT (1966) Stridulation of the garden tiger moth Arctia caja audible to the human ear. Proc R Entomol Soc Lond A 41:167–170

    Google Scholar 

  • Rowe C (1999) Receiver psychology and the evolution of multicomponent signals. Anim Behav 58:921–931

    Article  PubMed  Google Scholar 

  • Rowe C (2002) Sound improves visual discrimination learning in avian predators. Proc R Soc Lond B Biol Sci 269:1353–1357

    Article  PubMed  Google Scholar 

  • Rowe C, Guilford T (1996) Hidden colour aversions in domestic clicks triggered by pyrazine odours of insect warning displays. Nature 383:520–522

    Article  CAS  Google Scholar 

  • Rowe C, Guilford T (1999a) The evolution of multimodal warning displays. Evol Ecol 13:655–671

    Google Scholar 

  • Rowe C, Guilford T (1999b) Novelty effects in a multimodal warning signal. Anim Behav 57:341–346

    Article  PubMed  Google Scholar 

  • Rowe C, Guilford T (2001) The evolution of multimodal warning displays. Evol Ecol 13:655–671

    Google Scholar 

  • Ryan MJ (1980) Female mate choice in a neotropical frog. Science 209:523–525

    Google Scholar 

  • Ryan MJ (1985) Energetic efficiency of vocalization by the frog Physalaemus pustulosus. J Exp Biol 116:47–52

    Google Scholar 

  • Ryan MJ, Rand AS (1990) The sensory basis of sexual selection for complex calls in the Tungara frog, Physalaemus pustulosus (Sexual Selection for Sensory Exploitation). Evolution 44:305–314

    Google Scholar 

  • Ryan MJ, Wagner WE (1987) Asymmetries in mating preferences between species—female swordtails prefer heterospecific males. Science 236:595–597

    Google Scholar 

  • Savalli UM (1994) Tail length affects territory ownership in the yellow-shouldered widowbird. Anim Behav 48:105–111

    Article  Google Scholar 

  • Scheffer SJ, Uetz GW, Stratton GE (1996) Sexual selection, male morphology, and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae). Behav Ecol Sociobiol 38:17–23

    Article  Google Scholar 

  • Scheuber H, Jacot A, Brinkhof MWG (2003a) Condition dependence of a multicomponent sexual signal in the field cricket Gryllus campestris. Anim Behav 65:721–727

    Article  Google Scholar 

  • Scheuber H, Jacot A, Brinkhof MWG (2003b) The effect of past condition on a multicomponent sexual signal. Proc R Soc Lond B Biol Sci 270:1779–1784

    Article  PubMed  Google Scholar 

  • Schuler W, Roper TJ (1992) Responses to warning coloration in avian predators. Adv Stud Behav 21:111–146

    Google Scholar 

  • Smith C, Reynolds JD, Sutherland W, Jurajda P (2000) Adaptive host choice and avoidance of superparasitism in the spawning decisions of bitterling (Rhodeus sericeus). Behav Ecol Sociobiol 48:29–35

    Article  Google Scholar 

  • Smith JW (1965) Message, meaning, and context in ethology. Am Nat 97:405–409

    Article  Google Scholar 

  • Spence C, Nicholls MER, Gillespie N, Driver J (1998) Cross-modal links in exogenous covert spatial orienting between touch, audition, and vision. Percept Psychophys 60:544–557

    CAS  PubMed  Google Scholar 

  • Stokes D (1979) Stokes nature guides: a guide to bird behavior, vol I. Little, Brown, Boston

    Google Scholar 

  • Stokes D, Stokes L (1983) Stokes nature guides: a guide to bird behavior, vol II. Little, Brown, Boston

    Google Scholar 

  • Stokes D, Stokes L (1998) Stokes nature guides: a guide to bird behavior, vol III. Little, Brown, Boston

    Google Scholar 

  • Sumby WH, Pollack I (1954) Visual contribution to speech intelligibility in noise. J Acoust Soc Am 26:1298–1319

    Google Scholar 

  • Tibbetts EA (2002) Visual signals of individual identity in the wasp Polistes fuscatus. Proc R Soc Lond B Biol Sci 269:1423–1428

    Article  PubMed  Google Scholar 

  • Tregenza T, Wedell N (2000) Genetic compatibility, mate choice and patterns of parentage: invited review. Mol Ecol 9:1013–1027

    Article  CAS  PubMed  Google Scholar 

  • Uetz GW, Roberts JA (2002) Multisensory cues and multimodal communication in spiders: insights from video/audio playback studies. Brain Behav Evol 59:222–230

    Article  PubMed  Google Scholar 

  • Van Staaden MJ, Romer H (1997) Sexual signalling in bladder grasshoppers: tactical design for maximizing calling range. J Exp Biol 200:2597–2608

    PubMed  Google Scholar 

  • Von Helversen D (1984) Parallel processing in auditory pattern recognition and directional analysis by the grasshopper Chorthippus biguttulus L. (Acrididae). J Comp Physiol 154:837–846

    Google Scholar 

  • Wedekind C, Meyer P, Frischknecht M, Niggli UA, Pfander H (1998) Different carotenoids and potential information content of red coloration of male three-spined stickleback. J Chem Ecol 24:787–801

    Article  CAS  Google Scholar 

  • Wells KD (1977) The courtship of frogs. In: Taylor DH, Sheldon IG (eds) The reproductive biology of amphibians. Plenum, New York, pp 233–262

  • Wiley RH (1994) Errors, exaggeration, and deception in animal communication. In: Real L (ed) Behavioral mechanisms in evolutionary ecology. University of Chicago Press, Chicago

  • Wilson EO (1975) Sociobiology. The new synthesis. Harvard University Press, Cambridge

  • Zahavi A (1975) Mate selection—selection for a handicap. J Theor Biol 53:205–214

    CAS  PubMed  Google Scholar 

  • Zahavi A (1987) The theory of signal selection and some of its implications. In: Delfino V (ed) International symposium of biological evolution. Adriatica Editrice, Bari, pp 305–327

  • Zuk M, Ligon JD, Thornhill R (1993) Effects of experimental manipulation of male secondary sex characteristics on female mate preference in red jungle fowl. Anim Behav 44:999–1006

    Google Scholar 

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Acknowledgements

We would like to thank R. Hoy, K. Reeve, J. Bradbury, N. Vander Sal, D. Elias, and members of the Papaj Lab for commenting on earlier drafts of this manuscript. We would also like to thank A. Cutter and W. Maddison for fruitful discussions and two anonymous reviewers for invaluable suggestions and insights.

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Hebets, E.A., Papaj, D.R. Complex signal function: developing a framework of testable hypotheses. Behav Ecol Sociobiol 57, 197–214 (2005). https://doi.org/10.1007/s00265-004-0865-7

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