Community structure of mesopelagic fishes constituting sound scattering layers in the eastern tropical North Atlantic

Abstract

Mesopelagic sound scattering layers (SSLs) are predominantly associated with mesopelagic fish taxa with gas-inflated swimbladders that perform active nocturnal diel vertical migration (DVM), like species of the families Myctophidae, Gonostomatidae, Sternoptychidae and Phosichthyidae. Larger-sized species with limited, partial, asynchronous and non-migratory migration pattern (non-DVM) and without gas-bearing swimbladders are presumably invisible by hydroacoustic methods operating at lower frequencies (18–38 kHz). Their vertical migration behaviour and functional role in the mesopelagic community remain largely disregarded. The present study investigated (i) the taxonomic and functional mesopelagic fish composition (migration behaviour and feeding guild as traits) of the main SSLs and (ii) regional variation in the vertical community structure related to the presence of the oxygen minimum zone (OMZ) in the eastern tropical North Atlantic. We analysed hydroacoustic and biological trawl data collected concurrently with a pelagic midwater trawl in distinct SSLs between 45 and 680 m depth by means of ordination, cluster and compositional analyses. Whereas daytime hydroacoustic backscatter profiles demonstrated higher mean Sv (38 kHz) backscatter at shallower depths at low-oxygen (LO) station 309 compared to the equatorial (EQ) station 330, night-time hydroacoustic profiles at LO stations could not be directly related to OMZ conditions. Catch abundances and biomass showed no clear regional pattern between EQ and LO stations. The analysis of trawl samples emphasised the importance of non-DVM species to the composition of tropical SSLs and further indicated regional and vertical variation in the mesopelagic fish community structure. At all stations, non-DVM species made up the largest proportions of total tow community composition (abundance and biomass) in the principal and secondary mesopelagic SSL during night-time (375–680 m) and during most daytime tows (LO: 390–555 m, EQ: 325–500 m). Our analysis suggests a more pronounced gradient structure in the vertical community composition of the LO compared to the EQ. On the one hand, shallower tows at eastern LO stations 306 and 311 were clustered with comparatively deeper tows. The eastern LO stations 306–315 had the highest numbers of non-DVM species in the principal epi- and mesopelagic SSL and the largest proportions of mesopelagic predatory species in the principal mesopelagic SSL. On the other hand, only the dominant DVM-species of the LO (i.e., Nannobrachium isaacsi, Ceratoscopelus warmingii, Notoscopelus resplendens, Lepidophanes guentheri and Hygophum macrochir) also showed consistent presence in the principal mesopelagic SSL during night-time. Moreover, our study demonstrates the usefulness of using biomass as additional parameter, when investigating functional aspects of mesopelagic community structure.