Abondance et diversité acoustique des populations de poissons dans la baie de Calvi.

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Abondance et diversité acoustique des populations de poissons dans la baie de Calvi. 1 MORFONCT, Laboratoire de Morphologie Fonctionnelle et Evolutive, ULiège, Belgium. 2 CHORUS Institute, Grenoble, France. 3 STARESO Research Station, Calvi, France. Marta Bolgan 1, Lucia Di Iorio 2, Pierre Lejeune 3 & Eric Parmentier 1

Fish acoustic communication More than 800 species have evolved morphological, physiological and neurological adaptations allowing them to rely on sound for communication Fish sounds are especially conspicuous during the breeding season, in relation to agonistic interactions and mating activities -individual recognition, mate choice, readiness to spawn, social status assessment etc- Intra-specific variation of fish sounds is generally smaller than inter-specific variation. Once the sound has ben characterized, the stereotypical nature of fish sounds makes it relatively easy to identify which species of fish vocalize in the wild.

Using fish sounds as natural tags: Passive Acoustic Monitoring (PAM) in fishery science Non-invasive census Continuous monitoring Independent of weather, time of day and human effort Diel, seasonal & geographical pattern of activity Habitat preferences Pinpoint fish spawning grounds Evaluating the effects of man-made noise Conservation/ Management programs

Fish PAM: MORFONCT at STARESO Sound and sonic apparatus characterisation, location of vocal species in the wild Carapus acus (Parmentier et. al 2006) Epinephleus marginatus (Bertucci et et al. 2015) Ophidion rochei (Kéver et al. 2016) Year-round characterisation of fish vocal activity Ophidion rochei (Kéver et al. 2016) Influence of environmental conditions on fish vocal behaviour Ophidion rochei (Kéver et al. 2015) Consistency of fish call features along a Mediterranean gradient Sciaena umbra (Parmentier et al. 2017) Carapidae>Carapus acus Serranidae>Epinephleus marginatus Ophidiidae>Ophidion rochei Parmentier, E., Fine, M., Vandewalle, P., Ducamp, J. J., Lagardère, J. P. (2006). Acta Zoologica, 87(2), 113 119. Bertucci, F., Lejeune, P., Payrot, J., Parmentier, E. (2015). Journal of Fish Biology, 87(2), 400 421. Kéver, L., Lejeune, P., Michel, L. N., Parmentier, E. (2016). Marine Ecology, 37(6), 1315 1324. Kéver, L., Boyle, K. S., Parmentier, E. (2015). Journal of Fish Biology, 87(2), 502 509. Parmentier, E., Di Iorio, L., Picciulin, M., Malavasi, S., Lagardère, J. P., Bertucci, F. (2017). Animal Conservation doi:10.1111/acv.12362 Sciaenidae> Sciaena umbra

Acoustic community Aggregation of species that produce sounds by using internal or external soundproducing tools and which interact acoustically in a specific habitat (Farina & James, 2016). Fish acoustic communication has rarely been studied at community level (Ruppè et al. 2015).

Acoustic niche hypothesis Individuals in acoustic communities compete for the use of the sound resource for communication (niche competition) ACOUSTIC NICHE HYPOTHESIS (Krause, 1993) To avoid interference, fish species sharing the same acoustic space have co-evolved to exploit different frequency bands Frequency partition (Ruppè et al. 2015) And/ or to emit sounds at different time of the day/ year Temporal partition Ruppé, L., Clément, G., Herrel, A., Ballesta, L., Décamps, T., Kéver, L., & Parmentier, E. (2015). Environmental constraints drive the partitioning of the soundscape in fishes. Proceedings of the National Academy of Sciences, 112(19). doi.org/10.1073/pnas.1424667112

Testing the acoustic niche hypothesis in Mediterranean fish communities Abundance and diversity variation of fish sounds at different depths in STARESO (and comparison with other sites with similar environmental conditions) -20 m -40 m -125 m -150m -162 m Posidonia oceanica Sandy area Head of the canyon One month of simultaneous recordings in the peak of fish vocal season (July) in three Mediterranean meadows (Calvi bay, Mallorca, Crete) (2017) One year of recordings in the Calvi bay (Kéver et al. 2016) Analysis over one month (June 2013) Short-term SAM during August and October (2016-2107) in the Calvi bay (1 to 2 nights)

-20 m Posidonia oceanica Data analysis ongoing, preliminary results

Fish active acoustic space in P. oceanica at -20 m: one month of recording (743 hours: July) Day-time: Boat traffic Night-time: Fish calls

STARESO, -20 m from 5pm to 6am Most common fish sounds 1 1. Kwa Sound characterised in details by Di Iorio et al. (2018) Unknown emitter (but studies are ongoing) 20 times more abundant than the other fish sounds FREQUENCY PARTITION 3 2 2. Ophidion rochei male calls 3. Sciaena umbra calls TEMPORAL PARTITION (but niche overlap can occur)

-40 m Sandy area

STARESO, -40 m sandy area Analysis: one month (July) Sound diversity: five different types of fish sounds for only two, the specific identity of the emitter is known 1. Kwa = harmonic sound of unknown origin 2. S = Sciaena umbra sound 3. O = Ophidion rochei sound (male) 4. PS = pulse series sounds of unknown origin; 5. OP = single pulse of unknown origin. Variable PCA1 PCA2 Peak frequency (Hz) -0.444 0.688 Duration (s) 0.532 0.389 Pulse period (s) 0.519-0.343 Number of pulses 0.501 0.534

STARESO -40 m sandy area Kwa -20m Relative percentage Kwa -40m Time of the day Kwa at -40m less pseudo-harmonics Narrower band energy Weaker sound Habitat preference vs acoustic communities boundaries?

900 STARESO -40 m sandy area Peak freqeuncy (Hz) 800 700 600 500 400 300 200 Frequency 95% 5% 100 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 Hour of the day S= S. umbra sounds; O= O. rochei sounds; Kwa = harmonic sounds of unknown origin; PS = pulse series of unknown origin; OP = single pulse of unknown origin.

-125 to -162 m Head of the canyon October 2016, 31 hours (continuous) 04/10/2016 10:00 to 05/10/2016 17:08 SH = -155 db, G = 0 db, D = 2.5V Depth ~ 162 m Depth hydrophone ~ 157 m August 2017, 44 hours (continuous) 29/08/2017 14:20 to 31/08/2017 10:40 SH = -170.2 db, G = 4 db, D = 1.58 V Depth ~ 150 m Depth hydrophone ~ 142 m October 2017, 31 hours (continuous) 02/10/2017 09:00 to 04/10/2017 14:00 SH = -164 db, G = 10 db, D = 1V Depth ~ 125m Depth Hydrophone ~ 120 m MORFONCT + CHORUS DATA

STFRP SOUND DIEL PATTERN (n sound x min -1 ) Range= 0-12 sound x min -1-125 m October Ophidion sp. STFRP Range= 0-2 sound x min -1 Pulse series Single boom -150 m August The cloud (fish??) -162 m October

STFRP: Stereotyped Trains of Fast-Repeated Pulses October -125m -162 m

Train duration STFRP: Stereotyped Trains of Fast-Repeated Pulses Train period STFRP duration Pulse period Cycle duration

August -150 m Ophidion sp. 3 6.5 3 6.5 pulse period (ms) 180 160 140 120 100 80 60 40 20 0 1 3 5 7 9 11131517192123252729313335373941 pulse number

What happens during daytime hours? NIGHT, lower levels of man-made noise (30 August 2017, -150 m, 1:20 am) Man-made noise= Lo-Fi acoustic space DAY, increase of man-made noise (boat traffic) (30 August 2017, - 150 m, 7:11 am)

Fish acoustic communities in the Calvi bay PAM= has the potential of providing continuous, not-invasive monitoring of fish community dynamics over large spatial and temporal scales Coastal areas (-20, -40 m) during summer months Frequency partition (e.g. Kwa) and temporal partition with partial niche overlap (e.g. Sciaena umbra and Ophidion rochei) Differential frequency range and intensity of the same sound in different environments proxy for species richness/ habitat selections? Head of the canyon (-125, -150, -162) Fish sounds detected with a small sampling effort re-inforce the H0: fish species adapted to deeper habitats exploit acoustic communication Different sound types at different depths Same sound type at different rates proxy for habitat selection/ spawning habits? Preliminary results: More studies are needed! In all areas, during daytime -increase in man-made noise -impaired detection of fish sounds -Lo-Fi acoustic space Which strategies in Lo-Fi environment? e.g. more sounds, louder sounds (individual level)? Fish that do not have such an adaptive capacity might be most impacted by the presence of man-made noise Which effects on an evolutionary scale? No one knows.. Precautionary approach when managing man-made noise.

Thanks for your attention! Marta Bolgan, Ph.D. MORFONCT, Laboratoire de Morphologie Fonctionnelle et Evolutive, ULiège, Belgique marta.bolgan@gmail.com mbolgan@ulg.ac.be