The results presented last year did not satisfy the commission, which had "great difficulty in understanding in the information provided what was relevant to THEMISTO or MOBYDICK". In the first evaluation document, it was specified that the valorisation should only concern the THEMISTO 2016 and 2017 campaigns. However, after clarification with the committee, it is indeed the valorisation of the entire series that is concerned, from 2016 to 2019, thus also including the year 2018 when the THEMISTO and MOBYDICK programs were associated (moreover referred to as such by the DOI: 10.17600/18000403). In order to facilitate the understanding of the commission on the valorisation of this double configuration, the presentation of the results will be in two parts: the THEMISTO program alone from 2016 to 2019, and then the valorisation of the joined MOBYDICK-THEMISTO program/campaign.
The major results strictly related to the THEMISTO program 2016-2019 are presented first. Contrary to what was proposed last year on the presentation of the valorisation of the previous acoustic campaigns because of their coherence with THEMISTO, the results obtained recently on the acoustic data linked to the previous MyctO-3D-MAP program are no longer presented here. The results obtained in the 2018 MOBYDICK-THEMISTO program will then be presented, presenting only the studies carried out in relation to the intermediate trophic levels addressed by THEMISTO.
1. Main results of the THEMISTO campaigns 2016-2019
- Study of zooplankton abundance variations in the Indian part of the Southern Ocean: influence of nycthemeral migrations and environmental parameters (Master Claire Godet)
The Joint MDCPR/REPCCOAI and MyctO/THEMISTO campaigns (from 2013 to 2016) have quantified zooplankton densities using two methods: the "Continuous Plankton Recorder" (CPR), which samples at the surface, and acoustics, which allows the visualization of zooplankton densities on the water column. Generalized linear mixed models were performed in order to analyze whether the variations in abundance of zooplankton collected by CPR could be explained by the nychtemeral cycles observed using acoustics, and to analyze whether environmental parameters could explain these fluctuations in abundance on CPR samples at night, the time of day when zooplankton abundance at the surface is at a maximum. By coupling the CPR data with the acoustic data, the fluctuations in abundance could be explained by the nycthemeral migrations. The bioregionalization of the South Indian and the comparison with zooplankton communities studied by the CPR collection could thus be carried out (Fig. ? from Godet et al. 2020), as well as the ecoregionalization of euphausiids and myctophids conducted in the framework of the REPCCOAI program.
Fig. 3. Map of the six bioregions and position of the three main fronts (from north to south: subtropical, subantarctic and polar front). Bioregions are defined based on abiotic parameters (SST, EKE and bathymetry) and Chl a concentration. From Godet et al. 2020
- Spatial structuring of pelagic ecosystems in the South Indian Ocean and the Indian zone of the Southern Ocean (Master Camille Merland)
The aim was to study the spatial structuring of micronektonic organisms in the South Indian zone, and particularly the scales at which they structure, and to highlight possible similarities with the scales at which top predators concentrate their foraging activity. In a first step, a functional approach allowed to regionalize and identify the patterns of variability of the micronekton. The regionalization obtained is consistent with that previously presented by Béhagle et al. 2016. It allowed us to take an additional step by prioritizing the patterns of variability. In two out of three years, the day/night distinction is the factor that predominantly structures the data. Spatial variability of densities is the second parameter that explains the difference between acoustic profiles. In contrast, we show an inverse trend in the third year in which the effect of spatial variability dominates over the effect of day/night variability. Next, we compared the scales of horizontal structuring between micronekton (prey) densities and predator foraging from telemetry monitoring. Common scales of the order of 20 km were revealed, suggesting the importance of the (sub-)mesoscale (attributed to oceanic fronts). This first approach combining trophic links and environmental forcing complements the approaches usually used to define areas of high conservation concern and will be further developed to reinforce the results obtained.
- "Switching off the Sun" to observe the twilight zone spatial dynamics across Saint-Paul and New-Amsterdam Islands, Southern Indian Ocean (PhD candidate Lloyd Izard)
Information on micronekton (> 2 cm organisms) are globally scarce in the open ocean and its vertical and horizontal distribution in relation to oceanographic structures is poorly known. The complex biodiversity compositing micronektonic functional groups leads to even more challenging interpretations of their spatial dynamics. Active acoustics is a non-intrusive observation method allowing high-frequency monitoring of micronektonic organism in the water column. Advanced generations of echosounders emit simultaneously several acoustic signals (multi-frequency device), allowing to gain a more representative view of the micronektonic community. While data becomes more abundant and complex, it is crucial to develop statistical tools aiming to objectively extract key components of its variability. In this study, we developed a Multivariate Functional Data Analysis method to determine patterns in micronekton structures across Saint-Paul and New-Amsterdam economic exclusive zone, at the boundary between large oceanic domains. Our results show a clear latitudinal acoustic pattern in concordance with hydrological features and biological samples (THEMISTO and REPCCOAI programs) (Fig. 4). This study proposes an objective method to analyse the vertical variability and quantify temporal or spatial modes of variability in multivariate acoustic data. Such methods could be implemented in any regions and allow 3-D modelling of micronektonic structuring.
Fig. 4. Classification of 2016 acoustic profiles (MD206) overlaid on maps of (from left to right) chlorophyll a concentrations (mg.m-3), temperature (°C) and surface salinity.
Amongst a compilation of data in the framework of an international programme, the active acoustic data were obtained with an EK80 echo sounder on board the Marion Dufresne during the THEMISTO 2018 campaign. The transect extends over 7766 km in the southwest Indian Ocean over an area ranging from 25 to 52° South and 51 to 77° East. The data were recorded from 05 January 2018 to 01 February 2018. Poor quality data due to unfavourable weather conditions were removed from the data set. After cleaning the data, an echo-integration was applied on the water column from 20 to 1000 m depth with a thickness of 10 m and a path length of 1 km. Maps of the distribution of average densities were thus created for the daytime and nighttime data and for each frequency studied. Similarly, the evolution of these density distributions was studied as a function of longitude and latitude along the entire route.
The concept of MESOPP is the creation of a collaborative network and associated e-infrastructure (marine ecosystem information system) between European and Australian research teams/institutes with similar interests in the Southern Ocean and Antarctic, the functioning of its marine ecosystems and the rapid changes occurring with global warming and marine resource exploitation. The THEMISTO acoustic measurements integrated the MESOPP reference datasets comprising 67 transects of 38 kHz volume backscatter intensity measurements (Sv, dB re 1 m-1) collected by 6 vessels (research and fishing) (Fig. 5). The reference data sets also contain calibrated and post-processed data from 18 kHz of 3 transits, 70 kHz of 2 transits, 120 kHz of 4 transits and 200 kHz of 2 transits. These reference data sets are from three areas of the Southern Ocean (South Atlantic, South Indian and South Pacific).
Fig. 5. MESOPP transits of calibrated and post-process acoustic backscatter
Post-processing of the data was undertaken using a combination of in-house tools developed by CSIRO, BAS and/or IRD and used to facilitate data management, data subset identification and post-processing. These tools used software packages including Java, Matlab, Movies 3D and Echoview. A preliminary comparison indicated that two of the three post-processing methods used generated similar values of water column NASC integrated from 0 to 800 m. There were some differences between biases with depth, although all still low. SEAPODYM LMTL modelling is optimised using the ratio between layers and therefore this discrepancy in bias with depth may cause greater uncertainty when using data post-processed with different methods. As a result it is clear that to ensure good data quality and assurance processes, data processing techniques should be compared on common data sets. Estimates of uncertainty to highlight precision and accuracy of the measurements and whether those variances matter in models need to be assessed so that potential errors are identified, and new and emerging methods are evaluated, in order to recommend best practice.
From the database referenced on the MESOPP website (https://www.mesopp.eu/), including THEMISTO data, analyses are underway and articles are being evaluated for publication in high-citation journals. THEMISTO contributes to the effort of sharing data for (meta)analysis at regional (Austral) or global scale.
2. Main results of the MOBYDICK-THEMISTO campaign (2018)
- Macrozooplankton/micronekton diversity, content and active carbon flux in contrasting environments of the Kerguelen region
Given the dominant role of the Southern Ocean in the global carbon cycle, it is important to consider estimates of active carbon flux relative to the global carbon budget. It is fundamental to assess the contribution of the active biological community of zooplankton and micronekton on the biological carbon pump. Our study provides new information on the response of the macrozooplankton and micronekton (MM) community under contrasting production regimes (oligotrophic vs. biologically enriched zones) found in the naturally fertilized Kerguelen Plateau site. Using a combination of trawl sampling and acoustic measurements, we demonstrated a vertical three-layer system at both the HNLC and fertilized stations (Fig. 6). However, different communities and biomasses and complex patterns of vertical distribution of MM and associated carbon fluxes were identified. This study provides new information on the importance of migrating biomasses and size structure in the biological pump of Southern Ocean regions under the influence of complex topography and land mass effects.
Fig. 6. Acoustic profiles and biomasses corresponding to the different taxa for the 4 stations of the MOBYDICK-THEMISTO program. From Cotté et al. in revision
- Pelagic food web structure in high nutrient low chlorophyll (HNLC) and naturally iron fertilized waters in the Kerguelen Islands region, Southern Ocean
Simultaneous collection of WP2, WP3, trawl and acoustic data was used to study the diversity, abundance and biomass of pelagic communities, as well as ecosystem dynamics in different primary production regimes in the Kerguelen region.
We found that the pelagic macrozooplankton/micronekton communities differed under distinct production conditions despite a clearly visible vertical three-layer system. As a result, HNLC waters upstream of the Kerguelen shelf, dominated by pico and nanophytoplankton, had mesozooplankton/macrozooplankton communities that were > 0.6 trophic positions higher than on the naturally iron-enriched shelf that supports large summer diatom blooms (Fig. 7). This suggests that the efficiency of biomass accumulation in iron-enriched regions depends on the presence of large zooplanktonic grazers that can efficiently consume and assimilate diatoms, such as the crustaceans that were abundant on the shelf. Salps accounted for almost half of the micronekton biomass and the population dynamics of this abundant gelatinous organism responded to the contrasting production regimes of the area. Complex models of the vertical distribution of macrozooplankton/micronekton and estimates of carbon flux mediated by migratory myctophids helped to provide insight into the main components and mechanisms of active carbon export in the region and how these are modulated by the complex topography and land mass effects.
Fig. 7.
- Demography, population growth rates and vertical migrations of salps (Salpa thompsoni) on the Kerguelen Plateau.
S. thompsoni are important grazers in the Southern Ocean, and have been shown to perform diel vertical migration to 500 m. However, estimates of their size and stage specific migration patterns are lacking. This work is both novel and timely as it provides the first insights into S. thompsoni size and stage migration patterns in the Kerguelen plateau region. This study provides the first information (from trawls and acoustics) on the vertical distribution of blastozooids and oozoids across day and night; identifying that the smallest individuals are the strongers migrators, and that larger individuals may occur at a deeper daytime residence depth than previously considered (> 800 m).
This study also aimed to obtain first estimates of S. thompsoni population dynamics and growth rates on the Kerguelen Plateau (southern Indian Ocean) at contrasting hydrological stations on the Kerguelen Plateau. Environmental conditions were important in determining the density and development of S. thompsoni populations. Growth rates (0.5-7.0% d-1 ) were higher than previously reported on the Antarctic Peninsula (0.3-4.6% d-1 ), but lower than near the Antarctic Polar Front (APF; 3.7-20.7% d-1 ). Despite warm surface waters (4-5°C), low chlorophyll a concentrations prevented salp populations from growing as rapidly as populations near the APF. The Kerguelen Plateau region diverts a branch of warm water southwards towards Antarctica. Further studies on the population dynamics of S. thompsoni over several seasons are needed to fully understand their importance on the Kerguelen Plateau and their potential to invade to higher latitudes.
3. Contributions of the THEMISTO program
- The EU-IUCN BEST initiative aims to support projects for biodiversity and ecosystem services in overseas territories. The SEECTOR program (Significant Ecological areas for pElagic species and ConservaTion challenge in the sOutheRn Indian Ocean, PI C. Cotté) has focused on the sub-Antarctic zone of the Indian Ocean to identify trophic hotspots, i.e. where the highest densities of pelagic organisms of intermediate trophic levels and top predators occur. The aim is to provide (i) a better understanding of the prey-predator link and the environmental phenomena at their origin and (ii) to better delimit these hotspots according to their dynamical characteristics.
- Within the framework of Lloyd Izard PhD thesis obtained at SU and the collaboration with the French Southern and Antarctic Territories (TAAFs), MyctO/THEMISTO data are involved in the scientific support of the file of extension of the marine reserve in the St Paul - Amsterdam EEZ and announced in February 2022 (https://www.outre-mer.gouv.fr/emmanuel-macron-annonce-lextension-de-la-reserve-naturelle-des-terres-australes-francaises-la).
- Contribution to dissemination activities in the framework of THEMISTO:
THEMISTO program blog during the 2019 Obsaustral sea campaign: https://www.ipsl.fr/Actualites/A-decouvrir/Carnet-de-campagne-THEMISTO-campagne-oceanique-hydroacoustique-dans-l-ocean-indien
Participation to an exhibition, "MYCTOPHIDAE, voyage en eau profonde", organized at the Institut Universitaire Européen de la Mer (Brest), and to a journal, La Lanterne, during the Festival Ressac (Fig. 8)
Fig. 8: Illustration of the exhibition "MYCTOPHIDAE, voyage en eau profonde"
References (selection of references on the THEMISTO program aims from the list of all publications automatically associated by SGC with the THEMISTO and MOBYDICK-THEMISTO campaigns)
- Cotté Cedric, Ariza A., Berne Adrien, Habasque Jeremie, Lebourges-Dhaussy Anne, Roudaut Gildas, Espinasse B., Hunt B.P.V., Pakhomov E.A., Henschke N., Péron C., Conchon A., Koedooder C., Izard L., Cherel Y. (2022). Macrozooplankton and micronekton diversity and associated carbon vertical patterns and fluxes under distinct productive conditions around the Kerguelen Islands. Journal Of Marine Systems, 226, 103650 (18p.). https://doi.org/10.1016/j.jmarsys.2021.103650
- Hunt Brian P.V., Espinasse Boris, Pakhomov Evgeny A., Cherel Yves, Cotté Cedric, Delegrange Alice, Henschke Natasha (2021). Pelagic food web structure in high nutrient low chlorophyll (HNLC) and naturally iron fertilized waters in the Kerguelen Islands region, Southern Ocean. Journal Of Marine Systems, 224, 103625 (10p.). https://doi.org/10.1016/j.jmarsys.2021.103625
- Henschke Natasha, Cherel Yves, Cotté Cedric, Espinasse Boris, Hunt Brian P.V., Pakhomov Evgeny A. (2021). Size and stage specific patterns in Salpa thompsoni vertical migration. Journal Of Marine Systems, 222, 103587 (5p.). https://doi.org/10.1016/j.jmarsys.2021.103587
- Henschke Natasha, Blain Stéphane, Cherel Yves, Cotte Cedric, Espinasse Boris, Hunt Brian P.V., Pakhomov Evgeny A. (2021). Population demographics and growth rate of Salpa thompsoni on the Kerguelen Plateau. Journal Of Marine Systems, 214, 103489 (10p.). Publisher's official version : https://doi.org/10.1016/j.jmarsys.2020.103489 , Open Access version : https://archimer.ifremer.fr/doc/00659/77147/
- Godet Claire, Robuchon Marine, Leroy Boris, Cotté Cedric, Baudena Alberto, Da Silva Ophélie, Fabri-Ruiz Salomé, Lo Monaco Claire, Sergi Sara, Koubbi Philippe (2020). Matching zooplankton abundance and environment in the South Indian Ocean and Southern Ocean. Deep-sea Research Part I-oceanographic Research Papers, 163, 103347 (12p.). Publisher's official version : https://doi.org/10.1016/j.dsr.2020.103347 , Open Access version : https://archimer.ifremer.fr/doc/00643/75474/