Papers by Michael Greenfield
ENES Winter Symposium 2019 programme-final
11:40 Michael Greenfield Rhythm interaction in social groups: Selective attention and social netw... more 11:40 Michael Greenfield Rhythm interaction in social groups: Selective attention and social networks 12:00 Mathilde Massenet Are there individual differences in dog calls before two months of age? 12:20 David Reby Physiological and perceptual correlates of masculinity in children's voices 12:40 Arthur Guibard Analysis of propagation constraints on acoustic communication networks among mountain Galliformes 1:00 Thibaut Marin-Cudraz Rock ptarmigan spatio-temporal acoustic activity at Arsine’s glacier
HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific r... more HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific research documents, whether they are published or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers. L’archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d’enseignement et de recherche français ou étrangers, des laboratoires publics ou privés. Animal choruses emerge from receiver psychology OPEN Michael Greenfield, Yareli Esquer-Garrigos, Réjane Streiff, Virginie Party
Annals of the New York Academy of Sciences
The study of human language is progressively moving toward comparative and interactive frameworks... more The study of human language is progressively moving toward comparative and interactive frameworks, extending the concept of turn-taking to animal communication. While such an endeavor will help us understand the interactive origins of language, any theoretical account for cross-species turn-taking should consider three key points. First, animal turn-taking must incorporate biological studies on animal chorusing, namely how different species coordinate their signals over time. Second, while concepts employed in human communication and turn-taking, such as intentionality, are still debated in animal behavior, lower level mechanisms with clear neurobiological bases can explain much of animal interactive behavior. Third, social behavior, interactivity, and cooperation can be orthogonal, and the alternation of animal signals need not be cooperative. Considering turn-taking a subset of chorusing in the rhythmic dimension may avoid overinterpretation and enhance the comparability of future empirical work.
European Journal of Entomology, 2016
http://www.eje.cz free of null alleles and conforming to Hardy-Weinberg expectations) has proven ... more http://www.eje.cz free of null alleles and conforming to Hardy-Weinberg expectations) has proven challenging in E. diurnus. Indeed, a set of 16 loci had been reported for this species before our study (Hockham et al., 1999; Hamill et al., 2006). According to the authors themselves, these loci displayed strong heterozygote defi cit, and the incidence of null alleles was considerable for some of them. In a preliminary trial we tested 13 of these available 16 loci on samples from highly divergent populations previously characterized for mitochondrial DNA COI variation (Party et al., 2015). Most loci failed to amplify and/or presented complex allelic patterns impeding their scoring. This situation signifi cantly reduced the number of available markers to only fi ve, which is a minimum value for population genetic analyses. We therefore applied highthroughput (pyrosequencing) technology to a partial genomic library enriched in microsatellite motifs in order to increase the number of loci and fi lter out those of low quality according to criteria detailed below. MATERIAL AND METHODS Sample collection and DNA extraction Fifty-one specimens of E. diurnus were collected from nine localities in southern France between 2011 and 2014 (Fig. 1). Hind femora were dissected and preserved in 95% ethanol for DNA
Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the Tettigoniidae
Annals of the Entomological Society of America, 2003
OUR APPRECIATION OF biological diversity owesmuch to the collective efforts of a special cadre of... more OUR APPRECIATION OF biological diversity owesmuch to the collective efforts of a special cadre of scientists who become thoroughly captivated by particular organisms, recognize those central issues that can be studied with greatest precision in their favorite species, andenthusiastically advertise theirÞndings to the biological community. The Canadian behavioral ecologist Darryl Gwynne is among this breed, and his efforts have put katydids (bush-crickets, should you reside outside the Western Hemisphere) on the behavioral ecology and evolutionmap. True, bymid 20 century, katydids (Tettigoniidae) had already acquired some status in the natural history of North America through their mating songs, especially the variable syllables sung by Pterophylla camellifolia (true katydid). In addition, entomologists were becoming aware of the importance of these songs in species recognition, sexual advertisement, and male rivalry. But it was Darryl GwynneOs studies of mating systems inNorthAmerican andAustralian tettigoniids over the past 20 years Ð which have taught us so much about the evolution of courtship behavior and how ecological factors may insuence the roles assumed by the sexes Ð that earned this family a central place in biology. These landmark studies, accompanied by concise reviewsof theevolution, anatomy,physiology, and ecology of tettigoniids, are now presented in an eminently readable book, “Katydids and bush-crickets: reproductive behavior and evolution of the Tettigoniidae” (D.T. Gwynne, 2001, Cornell Univ. Press, 317 pp.). In nine chapters, Gwynne introduces us to the global diversity of the Tettigoniidae and its phylogenetic relationships to other Orthoptera, describes the basic biology of this family, and then focuses on Þve topics in reproductive behavior: pairing of the sexes as facilitated by acoustic and vibratory signals, courtship feeding and the evolution of spermatophore attachments, sexual selection, risks incurred by the signaler and receiver, and sexual roles in courtship. The last topic is given special attention, as some tettigoniids are noteworthy for variation and reversals in sexual roles. Thus, we learn about ardent and aggressive females, coy and discriminating males who donate as much or more material investment to their offspring as their female partners do, and the circumstances under which the insects may revert to habits more beÞtting their gender.Gwynnewrites inapersonal style, almost chatty in places, but he does not treat these observations as mere natural history oddities. Rather, he makes exemplary use of “the exception probes the rule” tactic to test and reÞne our understanding of the relationships between the sexes in animal mating systems. “Katydids and bush-crickets: reproductive behavior and evolution of the Tettigoniidae” is not, nor did its author intend it to be, an encyclopedic coverage of these insects. Readers desiring in-depth treatment of speciÞc subjects including tettigoniid systematics, acoustic behavior, and neurobiology may be better served by consulting the (slightly out-of-date) edited volume “The Tettigoniidae: biology, systematics and evolution”(W.J.Bailey andD.C.F.Rentz, [eds.], 1990, Springer-Verlag, 395 pp.). Instead, Darryl GwynneOs recent book guides the reader through the complex interplay among ecology, evolution, and behavior that has shapedcourtship interactions,mating systems, and sexual differences. Sexual selection theory is clearly a prevailing theme, and its nuances are carefully explained so that one may evaluate the various hypotheses derived from its application. Such application is particularly well developed for analyzing the evolution of the tettigoniid spermatophore, which is accounted forbyno fewer than10potential explanations based largely on sexual selection. Importantly, the explanation thatemerges fromcurrentanalysis asmost likely in general (ejaculate protection) is not that which the author favored in his initial research (paternal investment). As indicated above, “Katydids and bush-crickets” is not a book about mechanisms. Given its stated focus, this is Þne, but there are several places in which I would have preferred a stronger physiological or genetic approach: biomechanical and neurophysiological factors, in addition to ecological ones, do constrain the sorts of songs that katydids can sing and hear, and genetic aspects of signaling, mating preferences, and changes leading to speciation deserve more attention. Similarly, could better understanding of development and nutritional biochemistry provide new insight to the katydid spermatophore? The book includes a number of valuable comparisons with other insects, but several critical ones were not made. For example, gregarization in the decticine katydidAnabrus simplex (Mormon Cricket) is discussed, but there is scant mention of the “locust phenomenon” in Old World Acrididae. Could information on factors regulating locust populations and their movement help explain how…
Proceedings of the National Academy of Sciences of the United States of America, Nov 29, 2016
Small animals typically localize sound sources by means of complex internal connections and baffl... more Small animals typically localize sound sources by means of complex internal connections and baffles that effectively increase time or intensity differences between the two ears. However, some miniature acoustic species achieve directional hearing without such devices, indicating that other mechanisms have evolved. Using 3D laser vibrometry to measure tympanum deflection, we show that female lesser waxmoths (Achroia grisella) can orient toward the 100-kHz male song, because each ear functions independently as an asymmetric pressure gradient receiver that responds sharply to high-frequency sound arriving from an azimuth angle 30° contralateral to the animal's midline. We found that females presented with a song stimulus while running on a locomotion compensation sphere follow a trajectory 20°-40° to the left or right of the stimulus heading but not directly toward it, movement consistent with the tympanum deflections and suggestive of a monaural mechanism of auditory tracking. Mor...
Philosophical Transactions of the Royal Society B: Biological Sciences
This theme issue assembles current studies that ask how and why precise synchronization and relat... more This theme issue assembles current studies that ask how and why precise synchronization and related forms of rhythm interaction are expressed in a wide range of behaviour. The studies cover human activity, with an emphasis on music, and social behaviour, reproduction and communication in non-human animals. In most cases, the temporally aligned rhythms have short—from several seconds down to a fraction of a second—periods and are regulated by central nervous system pacemakers, but interactions involving rhythms that are 24 h or longer and originate in biological clocks also occur. Across this spectrum of activities, species and time scales, empirical work and modelling suggest that synchrony arises from a limited number of coupled-oscillator mechanisms with which individuals mutually entrain. Phylogenetic distribution of these common mechanisms points towards convergent evolution. Studies of animal communication indicate that many synchronous interactions between the signals of neigh...
Rhythm interaction in animal groups: selective attention in communication networks
Philosophical Transactions of the Royal Society B: Biological Sciences
Animals communicating interactively with conspecifics often time their broadcasts to avoid overla... more Animals communicating interactively with conspecifics often time their broadcasts to avoid overlapping interference, to emit leading, as opposed to following, signals or to synchronize their signalling rhythms. Each of these adjustments becomes more difficult as the number of interactants increases beyond a pair. Among acoustic species, insects and anurans generally deal with the problem of group signalling by means of ‘selective attention’ in which they focus on several close or conspicuous neighbours and ignore the rest. In these animals, where signalling and receiving are often dictated by sex, the process of selective attention in signallers may have a parallel counterpart in receivers, which also focus on close neighbours. In birds and mammals, local groups tend to be extended families or clans, and group signalling may entail complex timing mechanisms that allow for attention to all individuals. In general, the mechanisms that allow animals to communicate in groups appear to b...
Sexual selection goes dynamic
Peer Community In Evolutionary Biology
Evolution of synchronies in insect choruses
Biological Journal of the Linnean Society
Scientific Reports, 2016
Synchrony and alternation in large animal choruses are often viewed as adaptations by which coope... more Synchrony and alternation in large animal choruses are often viewed as adaptations by which cooperating males increase their attractiveness to females or evade predators. Alternatively, these seemingly composed productions may simply emerge by default from the receiver psychology of mate choice. This second, emergent property hypothesis has been inferred from findings that females in various acoustic species ignore male calls that follow a neighbor's by a brief interval, that males often adjust the timing of their call rhythm and reduce the incidence of ineffective, following calls, and from simulations modeling the collective outcome of male adjustments. However, the purported connection between male song timing and female preference has never been tested experimentally, and the emergent property hypothesis has remained speculative. Studying a distinctive katydid species genetically structured as isolated populations, we conducted a comparative phylogenetic analysis of the correlation between male call timing and female preference. We report that across 17 sampled populations male adjustments match the interval over which females prefer leading calls; moreover, this correlation holds after correction for phylogenetic signal. Our study is the first demonstration that male adjustments coevolved with female preferences and thereby confirms the critical link in the emergent property model of chorus evolution.
Ethology Ecology Evolution, May 19, 2010
A series of playback experiments conducted in a field arena showed that female tarbush grasshoppe... more A series of playback experiments conducted in a field arena showed that female tarbush grasshoppers, Ligurotettix planum (Orthoptera Acrididae), were attracted to male calls and that when given the choice of calls that differed only in relative timing, females oriented toward the leading calls. This psychoacoustic feature, known as a "precedence effect," occurred when a 0.2 or 1.0-sec silent interval separated the leading and following calls. Preference for leading calls disappeared at separations longer than 2 sec and when calls overlapped; in the latter situation, females even failed to exhibit phonotaxis. Previous work demonstrated that neighboring L. planum males time their calls in an alternating fashion and that they achieve this chorusing format with an "inhibitory resetting" mechanism that averts calling during the 2-sec interval following onset of a neighbor's call. We propose that time constants in this inhibitory resetting mechanism evolved under selection pressure from the female precedence effect. Intermale signal interactions occur in many acoustic orthopterans and anurans, and we predict that female precedence effects, presently known in only a few species, will be revealed as responsible for various of these interactions. As in L. planum and another orthopteran species (Neoconocephalus spiza) in which both male signal interactions and female psychoacoustic preferences have been studied, inhibitory intervals in male interactive calling are expected to be congruent with or to exceed the lengths of precedence effects.
Alem et al. 2014 What determines lek size?
Phenotypic plasticity and genotype x environment interactions in animal communication
Sexual activities in both males and females are expected to incur energetic costs and risk. We te... more Sexual activities in both males and females are expected to incur energetic costs and risk. We tested the expectation that females who exhibit a higher level of discrimination of potential mates' signals incur greater risk than females who are relatively indiscriminate by assaying mate choice and predator evasion in an acoustic moth, Achroia grisella. Female A. grisella evaluate males on the basis of their song, only orienting toward males whose ultrasonic songs are delivered at pulse rates above a lower threshold value. Non-flying females also respond negatively, by ceasing all movement, upon hearing ultrasonic pulses delivered at rates below an upper threshold value; this response is assumed to represent evasion of predatory bats, specifically gleaning species. We found that both the lower and upper pulse-rate thresholds varied considerably among individual females and that the upper pulse-rate threshold for defensive response was always equal to or slightly lower than the lower pulse-rate threshold for a mating response. Thus, females who exhibit heightened discrimination of potential mates' signals experience less, not more, predatory risk. This relationship may reflect the evolutionary origin of acoustic sexual communication in A. grisella as a coöption of the ancestral function of hearing in evasion of predatory bats.
Evolution of ultrasonic signalling in wax moths: discrimination of ultrasonic mating calls from bat echolocation signals and the exploitation of an antipredator receiver bias by sexual advertisement
Ethol Ecol Evol, 2000
Page 1. Evolution of ultrasonic signalling in wax moths: discrimination of ultrasonic mating call... more Page 1. Evolution of ultrasonic signalling in wax moths: discrimination of ultrasonic mating calls from bat echolocation signals and the exploitation of an anti-predator receiver bias by sexual advertisement MD GREENFIELD 1 and T. WEBER ... 1994) enhanc-ing bat avoidance. ...
sound waves in the near field propueirlt' in .r complex manner, ungoverned bv the phr 'ir'irl rul... more sound waves in the near field propueirlt' in .r complex manner, ungoverned bv the phr 'ir'irl rules applicable to the 'far field'. Sounrl prt's:rrrt' ancl particle velocity remain 90' or,rt-ol-phil'r' irt the near lield. P nrtic Iert elo c it11 re('(Jr [ () r's Animals may perceive sound \\'a\.r's l.r' tlr'r it'r'' sensitive to either particle velocitl-()r Irr-r'\\rlrr'. Particle-rrelocity deteCtorS fttr airtrontc \()ult(i t1'picalll' consist of sensory hairs lt'itlr rrssouiiilr'ti neuroltes. and they llnction as ntecltittto[t'r'r'ptors {Michelsen and Larserr, l9ti5 r. \\ hcn sitr-rated in the near field o1'relative 11' lon' , < I kHz) liequency sound waves, hairs orientcrl perpendicularly to the direction ol'rt'ave pr()prrgation are vibrated btt motiot't of'iltt'irir molecules, Amongst Orthoptera, nrechan()rrceptors on the cerci may operatc in tlris I'rrslriorr. These cercal structures fut-tction prirltrrrlr rn predator avoidance by cletecting the sirnplr prr'\ence of air movement ((}'ratzv uncl Ilrrstt'r'1. 1989), However, recent finclings slton tltirt some cercal mechanoreceptive cells in grr llids respoud to the 30-Hz 's1'llable' (pulst') rirle within conspecilic chirps (K:imper, I c.)fi5 t. Therelbre. these receptors ma1' be critical in close-ranse acoustic communicatior-t such as
Variation in Host-Plant Quality: Implications for Territoriality in a Desert Grasshopper
Ecology, Aug 1, 1987
... KELSEY R. DOWNUM3 Department of Ecology and Evolutionary Biology, University of California, I... more ... KELSEY R. DOWNUM3 Department of Ecology and Evolutionary Biology, University of California, Irvine, California 92717 USA Abstract. ... Ter-ritorial individuals stridulate during morning and mid-day hours and chase other males detected on the same shrub. ...
Ethology and Sociobiology, 1985
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Papers by Michael Greenfield