Holocentridae; acoustic communication; diversification; evolution; sonic apparatus; sound
Abstract :
[en] Holocentridae are nocturnal reef fishes that rely on acoustic communication in various behavioral contexts. Using a multidisciplinary approach combining phylogenetics, morphology, and acoustic data from 53 species, we reconstructed the evolutionary history of this communication system within the family. Our results reveal two main clades corresponding to the subfamilies Myripristinae and Holocentrinae, with congruent differences in both the morphology of the sound-producing apparatus and acoustic features. Interestingly, at finer taxonomic scales, closely related species often share similar sound-producing structures but can produce distinct sounds. This suggests that the evolution of acoustic communication in Holocentridae follows a three-step diversification pattern: (1) the emergence of structures enabling sound production; (2) differentiation of sound-producing mechanisms within each subfamily, likely constrained by internal factors such as the swim bladder morphology; and (3) further diversification through differentiated use of conserved mechanisms, consistent with a role of neurophysiological modulation. This scenario mirrors patterns observed in other functional traits, where morphological innovations open new functional pathways, followed by fine-scale diversification based on the flexible exploitation of similar structures. It provides one of the first evolutionary reconstructions of sound production mechanisms in fishes, expanding our understanding of the evolution of complex signaling systems.
Disciplines :
Zoology
Author, co-author :
Banse, Marine ; Université de Liège - ULiège > Département de Biologie, Ecologie et Evolution > Morphologie fonctionnelle et évolutive
Van Steenberge, Maarten; Operational Directorate Taxonomy and Phylogeny Royal Belgian Institute of Natural Sciences Brussels Belgium
Sonet, Gontran; Joint Experimental Molecular Unit (JEMU), Operational Directorate Taxonomy and Phylogeny Royal Belgian Institute of Natural Sciences Brussels Belgium
Magda, Denisa; Operational Directorate Taxonomy and Phylogeny Royal Belgian Institute of Natural Sciences Brussels Belgium
Akaike, H. 1974. “A New Look at the Statistical Model Identification.” IEEE Transactions on Automatic Control 19: 716–723.
Amorim, M. C. P., M. E. Knight, Y. Stratoudakis, and G. F. Turner. 2004. “Differences in Sounds Made by Courting Males of Three Closely Related Lake Malawi Cichlid Species.” Journal of Fish Biology 65: 1358–1371.
Amorim, M. C. P., and A. S. M. Neves. 2007. “Acoustic Signalling During Courtship in the Painted Goby, Pomatoschistus pictus.” Journal of the Marine Biological Association of the United Kingdom 87: 1017–1023.
Amorim, M. C. P., R. O. Vasconcelos, M. Bolgan, S. S. Pedroso, and P. J. Fonseca. 2018. “Acoustic Communication in Marine Shallow Waters: Testing the Acoustic Adaptive Hypothesis in Sand Gobies.” Journal of Experimental Biology 221: jeb.183681.
Andrews, J. V., J. P. Schein, and M. Friedman. 2023. “An Earliest Paleocene Squirrelfish (Teleostei: Beryciformes: Holocentroidea) and its Bearing on the Timescale of Holocentroid Evolution.” Journal of Systematic Palaeontology 21, no. 1: 2168571.
Banse, M., E. Bertimes, D. Lecchini, T. J. Donaldson, F. Bertucci, and E. Parmentier. 2024. “Sounds as Taxonomic Indicators in Holocentrid Fishes.” npj Biodiversity 3: 33.
Banse, M., N. Hanssen, J. Sabbe, et al. 2024. “Same Calls, Different Meanings: Acoustic Communication of Holocentridae.” PLoS One 19: 1–26.
Banse, M., D. Lecchini, J. Sabbe, et al. 2024. “Production of Sounds by Squirrelfish During Symbiotic Relationships With Cleaner Wrasses.” Scientific Reports 14: 11158.
Banse, M., L. Minier, D. Lecchini, and E. Parmentier. 2024. “Acoustic Mobbing Behaviour: Vocal Fish Responses to Predation Risk Through Sound Communication.” Marine Biology 171: 141.
Bass, A. H., and R. Baker. 1990. “Sexual Dimorphisms in the Vocal Control System of a Teleost Fish: Morphology of Physiologically Identified Neurons.” Journal of Neurobiology 21: 1155–1168.
Bass, A. H., M. A. Marchaterre, and R. Baker. 1994. “Vocal-Acoustic Pathways in a Teleost Fish.” Journal of Neuroscience 14: 4025–4039.
Bass, A. H., and J. R. McKibben. 2003. “Neural Mechanisms and Behaviors for Acoustic Communication in Teleost Fish.” Progress in Neurobiology 69: 1–26.
Bertucci, F., K. Maratrat, C. Berthe, et al. 2020. “Local Sonic Activity Reveals Potential Partitioning in a Coral Reef Fish Community.” Oecologia 193: 125–134.
Blomberg, S. P., T. Garland, and A. R. Ives. 2003. “Testing for Phylogenetic Signal in Comparative Data: Behavioral Traits Are More Labile.” Evolution 57: 717–745.
Borkin, L. Y., S. N. Litvinchuk, Y. M. Rozanov, and D. V. Skorinov. 2004. “On Cryptic Species (From the Example of Amphibians).” Entomological Review 84: S75–S98.
Bradbury, J. W., and S. L. Vehrencamp. 1998. Principles of Animal Communication. Sinauer Associates.
Braune, P., S. Schmidt, and E. Zimmermann. 2008. “Acoustic Divergence in the Communication of Cryptic Species of Nocturnal Primates (Microcebus ssp.).” BMC Biology 6: 19.
Briggs, J. C. 1974. Marine Zoogeography. McGraw-Hill Book Company.
Brownstein, C. D., R. C. Harrington, L. R. V. Alencar, et al. 2025. “Phylogenomics Establishes an Early Miocene Reconstruction of Reef Vertebrate Diversity.” Science Advances 11: eadu6149.
Burton, J. A., and A. Nietsch. 2010. “Geographical Variation in Duet Songs of Sulawesi Tarsiers: Evidence for New Cryptic Species in South and Southeast Sulawesi.” International Journal of Primatology 31: 1123–1146.
Carlson, B. A., and A. H. Bass. 2000. “Sonic/Vocal Motor Pathways in Squirrelfish (Teleostei, Holocentridae).” Brain, Behavior and Evolution 56: 14–28.
Casadevall, M., J. Matallanas, M. Carrasson, and M. Munõz. 1996. “Morphometric, Meristic and Anatomical Differences Between Ophidion barbatum L., 1758 and O. rochei Müller, 1845 (Pisces, Ophidiidae).” Publicaciones Especiales, Instituto Español de Oceanografía 21: 45–61.
Castelnau, F. D. 1875. “Researches on the Fishes of Australia.” Official Record. Philadelphia Centennial Exhibition. Part 7: 1–47.
Chernomor, O., A. von Haeseler, and B. Q. Minh. 2016. “Terrace Aware Data Structure for Phylogenomic Inference From Supermatrices.” Systematic Biology 65: 997–1008.
Coates, A. G., and J. A. Obando. 1996. “The Geologic Evolution of the Central American Isthmus.” In Evolution and Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, 21–56. University of Chicago Press.
Cole, J. A. 2008. “A New Cryptic Species of Cicada Resembling Tibicen dorsatus Revealed by Calling Song (Hemiptera: Auchenorrhyncha: Cicadidae).” Annals of the Entomological Society of America 101: 815–823.
Colleye, O., P. Vandewalle, D. Lanterbecq, D. Lecchini, and E. Parmentier. 2011. “Interspecific Variation of Calls in Clownfishes: Degree of Similarity in Closely Related Species.” BMC Evolutionary Biology 11: 365.
Coombs, S., and A. N. Popper. 1979. “Hearing Differences Among Hawaiian Squirrelfish (Family Holocentridae) Related to Differences in the Peripheral Auditory System.” Journal of Comparative Physiology 132: 203–207.
Couldridge, V. C. K., and G. J. Alexander. 2002. “Color Patterns and Species Recognition in Four Closely Related Species of Lake Malawi Cichlid.” Behavioral Ecology 13: 59–64.
Don, R. H., P. T. Cox, B. J. Wainwright, K. Baker, and J. S. Mattick. 1991. “‘Touchdown’ PCR to Circumvent Spurious Priming During Gene Amplification.” Nucleic Acids Research 19: 4008.
Dornburg, A., J. Moore, J. M. Beaulieu, R. I. Eytan, and T. J. Near. 2015. “The Impact of Shifts in Marine Biodiversity Hotspots on Patterns of Range Evolution: Evidence From the Holocentridae (Squirrelfishes and Soldierfishes).” Evolution 69: 146–161.
Dornburg, A., J. A. Moore, R. Webster, et al. 2012. “Molecular Phylogenetics of Squirrelfishes and Soldierfishes (Teleostei: Beryciformes: Holocentridae): Reconciling More Than 100 Years of Taxonomic Confusion.” Molecular Phylogenetics and Evolution 65: 727–738.
Drummond, A. J., and M. A. Suchard. 2010. “Bayesian Random Local Clocks, or One Rate to Rule Them All.” BMC Biology 8: 114.
Edgar, R. C. 2004. “MUSCLE: Multiple Sequence Alignment With High Accuracy and High Throughput.” Nucleic Acids Research 32: 1792–1797.
Elemans, C. P. H., J. H. Rasmussen, C. T. Herbst, et al. 2015. “Universal Mechanisms of Sound Production and Control in Birds and Mammals.” Nature Communications 6: 8978.
Felsenstein, J. 2004. Inferring Phylogenies. Sinauer Associates.
Fine, M. L., and E. Parmentier. 2015. “Mechanisms of Fish Sound Production.” In Sound Communication in Fishes, edited by F. Ladich, 77–126. Springer. https://doi.org/10.1007/978-3-7091-1846-7.
Fine, M. L., and E. Parmentier. 2022. “Fish Sound Production: The Swim Bladder.” Acoustics Today 18: 13.
Fricke, R. 2018. “Ostichthys spiniger, a New Species of Soldierfish From New Ireland, Papua New Guinea, Western Pacific Ocean (Teleostei: Holocentridae).” Ichthyological Research 65: 127–133.
Froese, R., and D. Pauly. 2024. “FishBase.” https://www.fishbase.org.
Gainer, H., K. Kusano, and R. F. Mathewson. 1965. “Electrophysiological and Mechanical Properties of Squirrelfish Sound-Producing Muscle.” Comparative Biochemistry and Physiology 14: 661–671.
Gerhardt, H. C., F. Huber, and A. M. Simmons. 2003. “Acoustic Communication in Insects and Anurans: Common Problems and Diverse Solutions.” Journal of the Acoustical Society of America 114: 559.
Goller, F., and R. A. Suthers. 1996. “Role of Syringeal Muscles in Gating Airflow and Sound Production in Singing Brown Thrashers.” Journal of Neurophysiology 75: 867–876.
Greenfield, D. W. 2002. “Holocentridae: Squirrelfishes (Soldierfishes).” In The Living Marine Resources of the Western Central Atlantic, edited by K. E. Carpenter, 1192–1202. Food and Agriculture Organization of the United Nations (FAO).
Greenfield, D. W., J. E. Randall, and P. N. Psomadakis. 2017. “A Review of the Soldierfish Genus Ostichthys (Beryciformes: Holocentridae), With Descriptions of Two New Species From Myanmar.” Journal of Ocean Science Foundation 26: 1–33.
Grzywacz, B., K. G. Heller, E. Warchałowska-śliwa, T. V. Karamysheva, and D. P. Chobanov. 2017. “Evolution and Systematics of Green Bush-Crickets (Orthoptera: Tettigoniidae: Tettigonia) in the Western Palaearctic: Testing Concordance Between Molecular, Acoustic, and Morphological Data.” Organisms, Diversity and Evolution 17: 213–228.
Hansen, T. F. 1997. “Stabilizing Selection and the Comparative Analysis of Adaptation.” Evolution 51: 1341–1351.
Heled, J., and A. J. Drummond. 2010. “Bayesian Inference of Species Trees From Multilocus Data.” Molecular Biology and Evolution 27: 570–580.
Henry, C. S. 1994. “Singing and Cryptic Speciation Insects.” Trends in Ecology & Evolution 9: 388–392.
Hoang, D. T., O. Chernomor, A. von Haeseler, B. Q. Minh, and L. S. Vinh. 2018. “UFBoot2: Improving the Ultrafast Bootstrap Approximation.” Molecular Biology and Evolution 35: 518–522.
Horch, K., and M. Salmon. 1973. “Adaptations to the Acoustic Environment by the Squirrelfishes Myripristis violaceus and M. pralinius.” Marine Behaviour and Physiology 2: 121–139.
Horvatić, S., L. Bem, S. Malavasi, et al. 2019. “Comparative Analysis of Sound Production Between the Bighead Goby Ponticola kessleri and the Round Goby Neogobius melanostomus: Implications for Phylogeny and Systematics.” Environmental Biology of Fishes 102: 727–739.
Horvatić, S., S. Malavasi, J. Vukić, et al. 2021. “Correlation Between Acoustic Divergence and Phylogenetic Distance in Soniferous European Gobiids (Gobiidae; Gobius Lineage).” PLoS One 16: e0260810.
Huelsenbeck, J. P., R. Nielsen, and J. P. Bollback. 2003. “Stochastic Mapping of Morphological Characters.” Systematic Biology 52: 131–158.
Huelsenbeck, J. P., and F. Ronquist. 2001. “MRBAYES: Bayesian Inference of Phylogenetic Trees.” Bioinformatics 17: 754–755.
Hunter, J. P. 1998. “Key Innovation and Ecology of Macroevolution.” Trends in Ecology & Evolution 13: 31–36.
Irwin, D. E., S. Bensch, and T. D. Price. 2001. “Speciation in a Ring.” Nature 409: 333–337.
Irwin, D. E., J. H. Irwin, and T. D. Price. 2001. “Ring Species as Bridges Between Microevolution and Speciation.” Genetica 112–113: 223–243.
Ivanova, N. V., T. S. Zemlak, R. H. Hanner, and P. D. N. Hebert. 2007. “Universal Primer Cocktails for Fish DNA Barcoding.” Molecular Ecology Notes 7: 544–548.
Johnsen, S. 2014. “Hide and Seek in the Open Sea: Pelagic Camouflage and Visual Countermeasures.” Annual Review of Marine Science 6: 369–392.
Jones, G. 1997. “Acoustic Signals and Speciation: The Roles of Natural and Sexual Selection in the Evolution of Cryptic Species.” Advances in the Study of Behaviour 26: 317–354.
Jordan, D. S., and B. W. Evermann. 1898. “The Fishes of North and Middle America.” Bulletin – United States National Museum 47: 2871.
Kotlyar, A. N. 2017. “Holocentridae From Borodino Submarine Elevation (Philippine Sea).” Journal of Ichthyology 57: 37–44.
Ladich, F. 2014. “Fish Bioacoustics.” Current Opinion in Neurobiology 28: 121–127.
Ladich, F., and A. N. Popper. 2004. “Parallel Evolution in Fish Hearing Organs.” In Evolution of the Vertebrate Auditory System, edited by G. A. Manley, R. R. Fay, and A. N. Popper. Springer. https://doi.org/10.1007/978-1-4419-8957-4.
Larouche, O., J. R. Hodge, L. R. V. Alencar, et al. 2020. “Do Key Innovations Unlock Diversification? A Case-Study on the Morphological and Ecological Impact of Pharyngognathy in Acanthomorph Fishes.” Current Zoology 66: 575–588.
Lin, H.-C., and P. A. Hastings. 2011. “Evolution of a Neotropical Marine Fish Lineage (Subfamily Chaenopsinae, Suborder Blennioidei) Based on Phylogenetic Analysis of Combined Molecular and Morphological Data.” Molecular Phylogenetics and Evolution 60: 236–248.
Looby, A., K. Cox, S. Bravo, et al. 2022. “A Quantitative Inventory of Global Soniferous Fish Diversity.” Reviews in Fish Biology and Fisheries 32: 581–595.
Lugli, M. 2010. “Sounds of Shallow Water Fishes Pitch Within the Quiet Window of the Habitat Ambient Noise.” Journal of Comparative Physiology A 196: 439–451.
Lugli, M., H. Y. Yan, and M. L. Fine. 2003. “Acoustic Communication in Two Freshwater Gobies: The Relationship Between Ambient Noise, Hearing Thresholds and Sound Spectrum.” Journal of Comparative Physiology A 189: 309–320.
Malavasi, S., S. Collatuzzo, and P. Torricelli. 2008. “Interspecific Variation of Acoustic Signals in Mediterranean Gobies (Perciformes, Gobiidae): Comparative Analysis and Evolutionary Outlook.” Biological Journal of the Linnean Society 93: 763–778.
Matsunuma, M., Y. Fukui, and H. Motomura. 2018. “Review of the Ostichthys japonicus Complex (Beryciformes: Holocentridae: Myripristinae) in the Northwestern Pacific Ocean, With Description of a New Species.” Ichthyological Research 65: 285–314.
Mayr, E. 1989. Histoire de La Biologie. Diversité, Évolution et Hérédité. Fayard.
Mélotte, G., X. Raick, R. Vigouroux, and E. Parmentier. 2019. “Origin and Evolution of Sound Production in Serrasalmidae.” Biological Journal of the Linnean Society 128: 403–414.
Mélotte, G., R. Vigouroux, C. Michel, and E. Parmentier. 2016. “Interspecific Variation of Warning Calls in Piranhas: A Comparative Analysis.” Scientific Reports 6: 36127.
Miller, M. A., W. Pfeiffer, and T. Schwartz. 2010. “Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Trees.” Proceedings of the Gateway Computing Environments Workshop (GCE), 1–8.
Moulton, J. M. 1958. “The Acoustical Behavior of Some Fishes in the Bimini Area.” Biological Bulletin 114: 357–374.
Muñoz, M. I., S. Goutte, J. Ellers, and W. Halfwerk. 2020. “Environmental and Morphological Constraints Interact to Drive the Evolution of Communication Signals in Frogs.” Journal of Evolutionary Biology 33: 1749–1757.
Myrberg, A. A., and R. J. Riggio. 1985. “Acoustically Mediated Individual Recognition by a Coral Reef Fish (Pomacentrus partitus).” Animal Behaviour 33: 411–416.
Myung, I. J. 2003. “Tutorial on Maximum Likelihood Estimation.” Journal of Mathematical Psychology 47: 90–100.
Nair, R. J., and S. Dineshkumar. 2016. “New Distributional Records of Three Soldier Fishes (Pisces: Holocentridae: Myripristis) From Indian Waters.” Marine Biodiversity Records 9: 89.
Near, T. J., and C. E. Thacker. 2024. “Phylogenetic Classification of Living and Fossil Ray-Finned Fishes (Actinopterygii).” Bulletin of the Peabody Museum of Natural History 65: 3–302.
Nelson, E. M. 1955. “The Morphology of the Swim Bladder and Auditory Bulla in the Holocentridae.” Fieldiana Zoology 37: 121–130.
Nelson, J. S. 2006. Fishes of the World. John Wiley & Sons Inc.
Nguyen, L., H. A. Schmidt, A. von Haeseler, and B. Q. Minh. 2015. “IQ-TREE: A Fast and Effective Stochastic Algorithm for Estimating Maximum-Likelihood Phylogenies.” Molecular Biology and Evolution 32: 268–274.
Onuki, A., and H. Somiya. 2007. “Innervation of Sonic Muscles in Teleosts: Occipital vs. Spinal Nerves.” Brain, Behavior and Evolution 69: 132–141.
Parmentier, E., R. Diogo, and M. L. Fine. 2017. “Multiple Exaptations Leading to Fish Sound Production.” Fish and Fisheries 18: 958–966.
Parmentier, E., S. Gaëlle, B. Renaud, et al. 2022. “Sound Production and Mechanism in the Cryptic Cusk-Eel Parophidion vassali.” Journal of Anatomy 241: 581–600.
Parmentier, E., J. P. Lagardère, Y. Chancerelle, D. Dufrane, and I. Eeckhaut. 2008. “Variations in Sound-Producing Mechanism in the Pearlfish Carapini (Carapidae).” Journal of Zoology 276: 266–275.
Parmentier, E., D. Lanterbecq, and I. Eeckhaut. 2016. “From Commensalism to Parasitism in Carapidae (Ophidiiformes): Heterochronic Modes of Development?” PeerJ 2016: 1–23.
Parmentier, E., D. Lecchini, B. Frederich, C. Brie, and D. Mann. 2009. “Sound Production in Four Damselfish (Dascyllus) Species: Phyletic Relationships?” Biological Journal of the Linnean Society 97: 928–940.
Parmentier, E., P. Vandewalle, C. Brié, L. Dinraths, and D. Lecchini. 2011. “Comparative Study on Sound Production in Different Holocentridae Species.” Frontiers in Zoology 8: 12.
Percy, D. M., G. S. Taylor, and M. Kennedy. 2006. “Psyllid Communication: Acoustic Diversity, Mate Recognition and Phylogenetic Signal.” Invertebrate Systematics 20: 431–445.
Picciulin, M., M. Bolgan, C. Facca, R. Fiorin, F. Riccato, and S. Malavasi. 2021. “Sound Discrimination of Two Sympatric, Threatened Fish Species Allows for Their in Situ Mapping.” Aquatic Conservation 31: 2103–2118.
Rabosky, D. L. 2017. “Phylogenetic Tests for Evolutionary Innovation: The Problematic Link Between Key Innovations and Exceptional Diversification.” Philosophical Transactions of the Royal Society, B: Biological Sciences 372: 20160417.
Raick, X., A. L. Godinho, G. Kurchevski, A. Huby, and E. Parmentier. 2023. “Bioacoustics Supports Genus Identification in Piranhas.” Journal of the Acoustical Society of America 154: 2203–2210.
Raick, X., A. Huby, G. Kurchevski, A. L. Godinho, and É. Parmentier. 2020. “Use of Bioacoustics in Species Identification: Piranhas From Genus Pygocentrus (Teleostei: Serrasalmidae) as a Case Study.” PLoS One 15: e0241316.
Raick, X., D. Lecchini, L. Kéver, O. Colleye, F. Bertucci, and É. Parmentier. 2018. “Sound Production Mechanism in Triggerfish (Balistidae): A Synapomorphy.” Journal of Experimental Biology 221, no. 1: jeb168948.
Rambaut, A., A. J. Drummond, D. Xie, G. Baele, and M. A. Suchard. 2018. “Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7.” Systematic Biology 67: 901–904.
Randall, J. E., and P. C. Heemstra. 1985. “A Review of the Squirrelfishes of the Subfamily Holocentrinae From the Western Indian Ocean and Red Sea.” Ichthyological Bulletin 49: 1–27.
Randall, J. E., and P. C. Heemstra. 1986. “Holocentridae.” In Smiths' Sea Fishes, edited by M. M. Smith and P. C. Heemstra, 415–427. Springer-Verlag-Smith Institute of Ichthyology.
Revell, L., and L. Harmon. 2022. Phylogenetic Comparative Methods in R. Princeton University Press.
Revell, L. J. 2024. “Phytools 2.0: An Updated R Ecosystem for Phylogenetic Comparative Methods (And Other Things).” PeerJ 12: e16505.
Reyes-Arias, J. D., B. Brady, E. A. Ramos, et al. 2023. “Vocalizations of Wild West Indian Manatee Vary Across Subspecies and Geographic Location.” Scientific Reports 13: 11028.
Rice, A. N., and A. H. Bass. 2009. “Novel Vocal Repertoire and Paired Swimbladders of the Three-Spined Toadfish, Batrachomoeus trispinosus: Insights Into the Diversity of the Batrachoididae.” Journal of Experimental Biology 212: 1377–1391.
Ronquist, F., M. Teslenko, P. van der Mark, et al. 2012. “MrBayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice Across a Large Model Space.” Systematic Biology 61: 539–542.
Ruppé, L., G. Clément, A. Herrel, et al. 2015. “Environmental Constraints Drive the Partitioning of the Soundscape in Fishes.” Proceedings of the National Academy of Sciences of the United States of America 112: 6092–6097.
Salmon, M. 1967. “Acoustical Behavior of the Mempachi, Myripristis berndti, in Hawaii.” Pacific Science 21: 364–381.
Spanier, E. 1979. “Aspects of Species Recognition by Sound in Four Species of Damselfishes, Genus Eupomacentrus (Pisces: Pomacentridae).” Zeitschrift für Tierpsychologie 51: 301–316.
Spezie, G., V. Torti, G. Bonadonna, et al. 2023. “Evidence for Acoustic Discrimination in Lemurs: A Playback Study on Wild Indris Indri indri.” Current Zoology 69: 41–49.
Stöver, B. C., and K. F. Müller. 2010. “TreeGraph 2: Combining and Visualizing Evidence From Different Phylogenetic Analyses.” BMC Bioinformatics 11: 7.
Tamura, K., G. Stecher, and S. Kumar. 2021. “MEGA11: Molecular Evolutionary Genetics Analysis Version 11.” Molecular Biology and Evolution 38: 3022–3027.
Trifinopoulos, J., L. Nguyen, A. von Haeseler, and B. Q. Minh. 2016. “W-IQ-TREE: A Fast Online Phylogenetic Tool for Maximum Likelihood Analysis.” Nucleic Acids Research 44: 232–235.
Uy, J. A. C., D. E. Irwin, and M. S. Webster. 2018. “Behavioral Isolation and Incipient Speciation in Birds.” Annual Review of Ecology, Evolution, and Systematics 49: 1–24.
Velásquez, N. A., J. Marambio, E. Brunetti, M. A. Méndez, R. A. Vásquez, and M. Penna. 2013. “Bioacoustic and Genetic Divergence in a Frog With a Wide Geographical Distribution.” Biological Journal of the Linnean Society 110: 142–155.
Warrant, E. J., and N. Adam Locket. 2004. “Vision in the Deep Sea.” Biological Reviews of the Cambridge Philosophical Society 79: 671–712.
Wiens, J. J. 1998. “Combining Data Sets With Different Phylogenetic Histories.” Systematic Biology 47: 568–581.
Wilkins, M. R., E. S. C. Scordato, G. A. Semenov, et al. 2018. “Global Song Divergence in Barn Swallows (Hirundo rustica): Exploring the Roles of Genetic, Geographical and Climatic Distance in Sympatry and Allopatry.” Biological Journal of the Linnean Society 123: 825–849.
Winn, H. E., and J. A. Marshall. 1963. “Sound-Producing Organ of the Squirrelfish, Holocentrus rufus.” Physiological Zoology 36: 34–44.
Winn, H. E., J. A. Marshall, and B. Hazlett. 1964. “Behavior, Diel Activities, and Stimuli That Elicit Sound Production and Reactions to Sounds in the Longspine Squirrelfish.” Copeia 1964: 413–425. https://doi.org/10.2307/1441036.
Yang, Z. 1994. “Maximum Likelihood Phylogenetic Estimation From DNA Sequences With Variable Rates Over Sites: Approximate Methods.” Journal of Molecular Evolution 39: 306–314.
Zelditch, M. L., and W. L. Fink. 1996. “Heterochrony and Heterotopy: Stability and Innovation in the Evolution of Form.” Paleobiology 22: 241–254.
