In between coral branches: describing the courtship behavior of the Mediterranean cardinalfish Apogon imberbis (Linnaeus, 1758)


Published: Sep 2, 2025
Keywords:
Reproductive behavior underwater photography marine animal forests mesophotic habitat use
CHIARA GREGORIN
MARTINA COPPARI
TEO MARROCCO
TORCUATO PULIDO MANTAS
CAMILLA ROVETA
TREVOR J. WILLIS
CARLO CERRANO
Abstract

The behavior of a pair of cardinalfish Apogon imberbis (Linnaeus, 1758) (Apogonidae) was investigated through the acquisition of time-lapse images at three-min-intervals over five consecutive days, encompassing both diurnal and nocturnal hours. We characterized the phases of courtship, quantified the frequency and alternation of activity patterns, and assessed their duration across the diel cycle, highlighting potential differences between sexes. The habitat preferences by A. imberbis associated to the red octocoral Paramuricea clavata (Risso, 1827) (Cnidaria, Anthozoa) were documented for the first time in the Mediterranean Sea. The pair exhibited pronounced diel behavioral patterns, with increased nocturnal activity likely linked to zooplankton foraging, and diurnal rest in shelters provided by the coral branches. The courtship consisted in three activities that were cyclically repeated throughout the day, with a total duration of 10.2 hours. The novel behavior named “patrolling” was played by the female after the mating event, presumably linked to the egg-laying site defense. The time-lapse technique offers the possibility to collect continuous, non-invasive observations at great depths and for prolonged periods, allowing for detailed surveys of the organisms, their interactions and the use of the habitat. The advantages and potentialities of this methodology could be further improved by machine learning programs for the recognition of animal activity patterns, allowing for the processing of massive dataset, reducing human workload and improving accuracy and time-efficiency.

Article Details
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  • Early View_vol. 26, n. 3, 2025
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References
Aguzzi, J., Fanelli, E., Ciuffardi, T., Schirone, A., De Leo, F.C. et al., 2018. Faunal activity rhythms influencing early community succession of an implanted whale carcass offshore Sagami Bay, Japan. Scientific Reports, 8 (1), 11163.
Azzola, A., Picchio, V., Asnaghi, V., Bianchi, C.N., Morri, C. et al., 2023. Troubles Never Come Alone: Outcome of Multiple Pressures on a Temperate Rocky Reef. Water, 15 (4), 825.
Barnett, A., Bellwood, D.R., 2005. Sexual dimorphism in the buccal cavity of paternal mouthbrooding cardinalfishes (Pisces: Apogonidae). Marine Biology, 148 (1), 205-212.
Bo, M., Costa, A., Coppari, M., Enrichetti, F., Bavestrello, G. et al., 2024. Diel rhythms of fish frequentation in a temperate mesophotic antipatharian forest and sleeping behavior of the red swallowtail perch Anthias anthias (Linnaeus, 1758). Marine Biology, 171 (9), 166.
Boero, F., Cicogna, F., Pessani, D., Pronzato, R., 1991. In situ observations on contraction behaviour and diel activity of Halcampoides purpurea var. mediterranea (Cnidaria, Anthozoa) in a marine cave. Marine Ecology, 12 (3), 185-192.
Bussotti, S., Guidetti, P., 2009. Do Mediterranean fish assemblages associated with marine caves and rocky cliffs differ? Estuarine, Coastal and Shelf Science, 81 (1), 65-73.
Bussotti, S., Guidetti, P., Belmonte, G., 2003. Distribution patterns of the cardinal fish, Apogon imberbis , in shallow marine caves in southern Apulia (SE Italy). Italian Journal of Zoology, 70 (2), 153-157.
Bussotti, S., Franco, A.D., Pey, A., Vieux-Ingrassia, J.-V., Planes, S. et al., 2017. Distribution patterns of marine cave fishes and the potential role of the cardinal fish Apogon imberbis (Linnaeus, 1758) for cave ecosystem functioning in the western Mediterranean. Aquatic Living Resources, 30, 15.
Cerrano, C., Danovaro, R., Gambi, C., Pusceddu, A., Riva, A. et al., 2010. Gold coral (Savalia savaglia) and gorgonian forests enhance benthic biodiversity and ecosystem functioning in the mesophotic zone. Biodiversity and conservation, 19 (1), 153-167.
Chabanet, P., Loiseau, N., Join, J.L., Ponton, D., 2012. VideoSolo, an autonomous video system for high-frequency monitoring of aquatic biota, applied to coral reef fishes in the Glorioso Islands (SWIO). Journal of Experimental Marine Biology and Ecology, 430, 10-16.
Chave, E.H., 1978. General Ecology of Six Species of Hawaiian Cardinalfishes. Pacific Science, 32 (3), 245-270.
De Brauwer, M., Gordon, L.M., Shalders, T.C., Saunders, B. J., Archer, M. et al., 2019. Behavioural and pathomorphological impacts of flash photography on benthic fishes. Scientific Reports, 9 (1), 748.
Del Rio, J., Aguzzi, J., Hidalgo, A., Bghiel, I., Manuel, A. et al., 2013. Citizen Science and marine community moni toring by video-cabled observatories: the OBSEA Citizen Science Project. p. 1-13. In: 2013 IEEE International Underwater Technology Symposium (UT), 5-8 March 2013, Tokyo, Japan.
Doya, C., Aguzzi, J., Pardo, M., Matabos, M., Company, J.B. et al., 2014. Diel behavioral rhythms in sablefish (Anoplopoma fimbria) and other benthic species, as recorded by the Deep-sea cabled observatories in Barkley canyon (NEPTUNE- Canada). Journal of Marine Systems, 130, 69-78.
Dunbrack, R., Zielinski, R., 2003. Seasonal and diurnal activity of sixgill sharks (Hexanchus griseus) on a shallow water reef in the Strait of Georgia, British Columbia. Canadian Journal of Zoology, 81 (6), 1107-1111.
Elhorst, R., Syposz, M., Wojczulanis-Jakubas, K., 2025. BEHAVE- facilitating behaviour coding from videos with AI-detected animals. Ecological informatics, 87, 103106.
Enrichetti, F., Baldrighi, E., Bavestrello, G., Betti, F., Canese, S. et al., 2022. Ecological role and phylogenetic position of a new habitat-forming species (Canalipalpata, Sabellidae) from the Mediterranean mesophotic soft bottoms. Estuarine, Coastal and Shelf Science, 265, 107737.
Fanelli, E., Masiá, P., Premici, A., Volpato, E., Da Ros, Z. et al., 2024. The re-use of offshore platforms as ecological observatories. Marine pollution bulletin, 209, 117262.
Francescangeli, M., Carandell, M., Mihai, T.D., Martínez, E., Nogueras, M. et al., 2023. Resource pulse in shallow waters: characterization of the scavenger community associated with a dolphin carcass. P. 1-5. In: OCEANS 2023-Limerick, 5-8 June 2023, Limerick, Ireland. Fukumori, K., Okuda, N., Yanagisawa, Y., 2009.
Female territoriality in a paternal mouthbrooding cardinalfish to avoid predation against spawned eggs. Canadian Journal of Zoology, 87 (6), 508-514.
Gardiner, N.M., Jones, G.P., 2005. Habitat specialisation and overlap in a guild of coral reef cardinalfishes (Apogonidae). Marine Ecology Progress Series, 305, 163-175.
Gardiner, N.M., Jones, G.P., 2010. Synergistic effects of habitat preference and gregarious behavior on habitat use in coral reef cardinalfish. Coral Reefs, 29 (4), 845-856.
Gori, A., Bavestrello, G., Grinyó, J., Dominguez-Carrió, C., Ambroso, S. et al., 2017. Animal Forests in Deep Coastal Bottoms and Continental Shelf of the Mediterranean Sea. Pp. 1-27. In: Marine Animal Forests. Rossi, S., Bramanti, L., Gori, A., Orejas, C. (Eds). Springer International Publishing, Cham.
Harmelin-Vivien, M.L., Harmelin, J.G., Chauvet, C., Duval, C., Galzin, R. et al., 1985. Evaluation visuelle des peuplements et populations de poissons méthodes et problèmes. Annual Review of Ecology, Evolution, and Systematics, 40 (4), 467-539.
Jacobsen, N.L., Roveta, C., Pulido Mantas, T., Coppari, M., Di Camillo, C.G. et al., 2024. Assessment of octocoral‐dominated benthic assemblages along a mesophotic gradient, with a focus on the impact of lost fishing gears. Aquatic Conservation: Marine and Freshwater Ecosystems, 34 (9), e4248.
Jeffreys, R.M., Lavaleye, M.S., Bergman, M.J., Duineveld, G.C., Witbaard, R., et al., 2010. Deep-sea macrourid fishes scavenge on plant material: evidence from in situ observations. Deep Sea Research Part I: Oceanographic Research Papers, 57 (4), 621-627.
Jeffreys, R.M., Lavaleye, M.S., Bergman, M.J., Duineveld, G.C., Witbaard, R. et al., 2011. Do abyssal scavengers use phytodetritus as a food resource? Video and biochemical evidence from the Atlantic and Mediterranean. Deep Sea Research Part I: Oceanographic Research Papers, 58 (4), 415-428.
Kerry, J.T., Bellwood, D.R., 2012. The effect of coral morphology on shelter selection by coral reef fishes. Coral Reefs, 31 (2), 415-424. Kolm, N., Berglund, A., 2004. Sex-specific territorial behaviour in the Banggai cardinalfish, Pterapogon kaunderni. Environmental Biology of Fishes, 70 (4) 375-379.
Kuwamura, T., 1983. Spawning Behavior and Timing of Fertilization in the Mouthbrooding Cardinalfish Apogon notatus. The Japanese Journal of Ichthyology, 30 (1), 61-71.
Kuwamura, T., 1985, Social and reproductive behavior of three mouthbrooding cardinalfishes, Apogon doederleini, A. niger and A. notatus. Environmental Biology of Fishes, 13 (1), 17-24.
Lehner, P.N., 1992. Sampling Methods in Behavior Research. Poultry Science, 71 (4), 643-649.
Linares, C., Coma, R., Garrabou, J., Díaz, D., Zabala, M., 2008. Size distribution, density and disturbance in two Mediterranean gorgonians: Paramuricea clavata and Eunicella singularis. Journal of Applied Ecology, 45 (2), 688-699.
Lu, A., Li, X.-Y., Lin, Y.-Y., Pan, Y.-J., 2024. Spawning, larval morphological ontogeny and first feeding of yellow-striped cardinalfish Ostorhinchus cyanosoma (Bleeker, 1853). Aquaculture, 583, 740622.
Luehrmann, M., Cortesi, F., Cheney, K.L., Busserolles, F., de Marshall, N.J., 2020. Microhabitat partitioning correlates with opsin gene expression in coral reef cardinalfishes (Apogonidae). Functional Ecology, 34 (5), 1041-1052.
Marnane, M.J., 2000. Site fidelity and homing behavior in coral reef cardinalfishes. Journal of Fish Biology, 57 (6), 1590-1600.
Mazzoldi, C., Randieri, A., Mollica, E., Rasotto, M.B., 2008. Notes on the reproduction of the cardinalfish Apogon imberbis form Lachea Island, Central Mediterranean, Sicily, Italy. Vie et Milieu/Life & Environment, 63-66.
Ménard, A., Turgeon, K., Roche, D.G., Binning, S.A., Kramer, D.L., 2012. Shelters and Their Use by Fishes on Fringing Coral Reefs. PloS ONE, 7 (6), e38450.
Montalbetti, E., Cavallo, S., Azzola, A., Montano, S., Galli, P. et al., 2023. Mucilage-induced necrosis reveals cellular oxidative stress in the Mediterranean gorgonian Paramuricea clavata. Journal of Experimental Marine Biology and Ecology, 559, 151839.
Okuda, N., 2001. The costs of reproduction to males and females of a paternal mouthbrooding cardinalfish Apogon notatus. Journal of Fish Biology, 58 (3) , 776-787.
Okuda, N., Yanagisawa, Y., 1996. Filial cannibalism by mouthbrooding males of the cardinal fish, Apogon doederleini, in relation to their physical condition. Environmental Biology of Fishes, 45 (4), 397-404.
Pierdomenico, M., Pandolfi, F., Mancini, G., Ventura, D., Falco, M. et al., 2024. Terrain, oceanographic, and biological factors underlying the development of Mediterranean coastal animal forests. Marine Environmental Research, 200, 106663.
Poliseno, A., Altuna, A., Cerrano, C., Wörheide, G., Vargas, S., 2017. Historical biogeography and mitogenomics of two endemic Mediterranean gorgonians (Holaxonia, Plexauridae). Organisms Diversity and Evolution, 17 (2), 365-373.
Raventos, N., 2007. Age, growth and reproductive parameters of the Mediterranean cardinal fish, Apogon imberbis. Journal of Applied Ichthyology, 23 (6), 675-678.
Rossi, S., Bramanti, L., Gori, A., Orejas, C., 2017. An Overview of the Animal Forests of the World. Marine Animal Forests, p.1-26.
Salvati, E., Provenzani, C., D’Ambrosi, A., Finoia, M.G., Romano, E. et al., 2023. In situ observation of circadian rhythm of polyps’ opening and closing of the coral Dendrophyllia ramea (Linnaeus, 1758). Mediterranean Marine Science, 24 (3), 574-585.
Saravanan, R., Vijayanand, P., Vagelli, A.A., Murugan, A., Shanker, S. et al., 2013. Breeding and rearing of the two striped cardinalfish, Apogon quadrifasciatus (Cuvier, 1828) in captive condition. Animal Reproduction Science, 137 (3- 4), 237-244.
Takeyama, T., Okuda, N., Yanagisawa, Y., 2007. Filial cannibalism as a conditional strategy in males of a paternal mouthbrooding fish. Evolutionary Ecology, 21 (1), 109-119.
Usuki, H., 1977. Underwater observation and experiments on pair formation and related behaviors of the Apogonid fish, Apogon notatus (Houttuyn). Contribution to Seto Marine Biology Laboratory, 24 (1-3), 223-243.
Vagelli, A.A., 1999. The Reproductive Biology and Early Ontogeny of the Mouthbrooding Banggai Cardinalfish, Pterapogon Kauderni (Perciformes, Apogonidae). Environmental Biology of Fishes, 56 (1), 79-92.
Vagelli, A.A., 2011. The Banggai Cardinalfish: Natural History, Conservation, and Culture of Pterapogon kauderni. New Jersey Academy for Aquatic Science, John Wiley & Sons (Eds), Wiley-Blackwell.
Vagelli, A.A., Volpedo, A.V., 2004. Reproductive Ecology of Pterapogon kauderni, an Endemic Apogonid from Indonesia with Direct Development. Environmental Biology of Fishes, 70 (3), 235-245.
Valletta, J.J., Torney, C., Kings, M., Thornton, A., Madden, J., 2017. Applications of machine learning in animal behaviour studies. Animal Behaviour, 124, 203-220.
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