Diversity, Composition, and Activities of Nano- and Pico-Eukaryotes in the Northern South China Sea With Influences of Kuroshio Intrusion

Nano- and pico-eukaryotes play important roles in the diversity and functions of marine ecosystems. Warm, saline, and nutrient-depleted water that originates in the Kuroshio Current seasonally intrudes into the northern South China Sea (NSCS) from autumn to spring. To clarify the mechanisms in shaping the community structure of nano- and pico-eukaryotes as well as impacts of the Kuroshio intrusion on the NSCS ecosystem, genomic DNA and RNA were co-extracted from samples collected at two depths from nine stations, and then the V9 region of 18S rDNA and rRNA was sequenced with high-throughput sequencing. Our results showed that Dinophyceae was the most diverse and abundant nanoeukaryotic group during the study period revealed by both DNA and RNA surveys. In contrast, the relative read abundance of MAST, Pelagophyceae, and Dinophyceae in the size fraction of picoeukaryotes might be largely underestimated by the DNA survey. The RNA survey was the more reliable method to investigate the eukaryotic community structure. Environmental filtering played an important role in shaping the community structure, and the sampling depth became the governing factor of the beta diversity under the environmental setting of stratification during the study period. The spatial variations in the diversity of nanoeukaryotes were subject to the dispersal limitation under the size rule. The effects of the Kuroshio intrusion on the nanoeukaryotic community structure might also be explained by the dispersal limitation. Overall, neutral processes are critical in shaping the community structure of nanoeukaryotes. The relative metabolic activities of nanoeukaryotes were relatively stable in accordance with the high similarity of community structure between sampling sites. The responses of the relative metabolic activities of picoeukaryotes to environmental factors displayed two distinct patterns: positive correlations with salinity and nutrients and negative with temperature for Dinophyceae, MAST, and Pelagophyceae, while reversed patterns for Mamiellophyceae and Radiolaria. Our findings improve the understanding of the nano- and pico-eukaryotic communities in the NSCS and the mechanisms of their assembly.

A Strong Kuroshio Intrusion into the South China Sea and Its Accompanying Cold-Core Anticyclonic Eddy in Winter 2020–2021

Multiple remote sensing datasets, combined with in-situ drifter observations, were used to analyze the Kuroshio intrusion through the Luzon Strait (LS). The results showed that a strong Kuroshio Current Loop (KCL) and accompanying anticyclonic eddy (ACE) existed in winter 2020–2021. As quantitatively demonstrated by the Double Index (DI), the Kuroshio Warm Eddy Index (KWI) had low values during a long sustained period compared to those in all other years in the available historical records. Remarkable kinematic properties (i.e., amplitude, diameter, propagated distance, lifespan and propagating speed) of the accompanying ACE were extracted by automatic eddy detection algorithms, showing that the ACE had a maximum diameter of 381 km and a peak amplitude of 50 cm, which significantly exceeded the previous statistics in winter. The orographic negative wind stress curl southwest of Taiwan Island and the westward Ekman transport through the LS during the winter half year of 2020–2021 both had large values beyond their historical maxima. Hence wind forcing is regarded as the primary forcing mechanism during this event. Alternating cyclonic eddies (CEs) and ACEs approaching on the east of the LS were identified, indicating that the interaction between the Kuroshio and the impinging CEs at proper locations made extra contributions to enhancing the KCL. The accompanying ACE had a distinctive feature of a cold-core structure at the surface layer, so as to be categorized as a cold core ACE (CC-ACE), and the temperature difference between the cold core and outer warm ring was maintained for three months. The generation and long duration of the CC-ACE may be due to the sustaining entrainment supported by the warm water from Kuroshio intrusion and the Northwest Luzon Coastal Current (NWLCC) successively.

Seasonal Variation of the Surface Kuroshio Intrusion into the South China Sea Evidenced by Satellite Geostrophic Streamlines

The long-term satellite altimeter and reanalysis data show that large seasonal variations are associated with geostrophic Kuroshio intrusion, but not with the current intensity, width and axis position east of Philippine. To address this issue, we examine the seasonal variability of surface intrusion patterns by a new streamline-based method. The along-streamline analysis reveals that the seasonality of geostrophic intrusion is only attributed to the cyclonic shear part of the flow, while the anticyclonic shear part always leaps across the Luzon Strait. A possible physical mechanism is proposed to accommodate these seasonal characteristics based on globally the vorticity (torque work) balance between the basin-wide negative wind stress curl and the positive vorticity fluxes induced by the lateral wall, as well as locally loss of balance between the torques of frictional stresses and normal stresses owing to the boundary gap. Through modifying the nearshore sea surface level, the northeasterly/southeasterly monsoon increases/decreases the positive vorticity fluxes in response to global vorticity balance, and simultaneously amplifies/alleviates the local imbalance by enhancing/reducing the positive frictional stress torque within the cyclonic shear layer. Therefore, in winter when the positive torque is large enough, the Kuroshio splits and the intrusion occurs, while in summer the stress torque is so weak that the entire current keeps flowing north.

Biogeographic Role of the Kuroshio Current Intrusion in the Microzooplankton Community in the Boundary Zone of the Northern South China Sea

Kuroshio Current intrusion (KCI) has significant impacts on the oceanographic conditions and ecological processes of the Pacific-Asian marginal seas. Little is known to which extent and how, specifically, the microzooplankton community can be influenced through the intrusion. Here, we focused on ciliates that often dominated the microzooplankton community and investigated their communities using high-throughput sequencing of 18S rRNA gene transcripts in the northern South China Sea (NSCS), where the Kuroshio Current (KC) intrudes frequently. We first applied an isopycnal mixing model to assess the fractional contribution of the KC to the NSCS. The ciliate community presented a provincial distribution pattern corresponding to more and less Kuroshio-influenced stations. Structural equation modeling revealed a significant impact of the KCI on the community, while environmental variables had a marginal impact. KCI-sensitive OTUs were taxonomically diverse but mainly belonged to classes Spirotrichea and Phyllopharyngea, suggesting the existence of core ciliates responding to the KCI. KCI-sensitive OTUs were grouped into two network modules that showed contrasting abundance behavior with the KC fraction gradient, reflecting differential niches (i.e., winner and loser) in the ciliate community during the Kuroshio intrusion scenarios. Our study showed that the Kuroshio intrusion, rather than environmental control, was particularly detrimental to the oligotrophic microzooplankton community.

Eddy-induced Heat Transport in the South China Sea

Eddy-induced heat transport (EHT) in the South China Sea (SCS) is important for the heat budget. However, knowledge of its variability is limited owing to discrepancies arising from the limitation of the down-gradient method and uncertainties arising from numerical models. Herein, we investigated the spatiotemporal variability and dynamics of EHT using a well-validated assimilated model. In particular, to the southeast of Vietnam (SEV) and west of Luzon Strait (WLS), significant values of annual mean EHT are observed and most EHT is confined in the upper 400 m. EHT also exhibits significant seasonality, and the largest EHT amplitude in autumn at SEV is mainly driven by the wind stress curl, while that in winter at WLS is mainly related to the Kuroshio intrusion. Energy budget analysis reveals that both the barotropic and baroclinic instabilities increase the eddy kinetic energy in autumn at SEV, whereas only the barotropic instability contributes to the eddy kinetic energy at WLS in winter. Specially, an up-gradient EHT is observed at WLS in all four seasons, characterized by the same directions between EHT and mean temperature gradient. The up-gradient EHT at WLS is induced by the baroclinic instability through an inverse energy transfer, which is generated by the interaction between the Kuroshio intrusion and topography below the surface layer. Moreover, the most significant up-gradient EHT in winter shows a wave-like southwestward propagating pattern in the subsurface layer.

Impacts of the Kuroshio Intrusion through the Luzon Strait on the Local Precipitation Anomaly

The Kuroshio Current has its origin in the northwestern Pacific, flowing northward to the east of Taiwan and the northern part of Luzon Island. As the Kuroshio Current flows northward, it quasi-periodically intrudes (hereafter referred to as Kuroshio intrusion (KI)) into the northern South China Sea (SCS) basin through the Luzon Strait. Despite the complex generation mechanisms of KI, the purpose of this study is to improve our understanding of the effects of KI through the Luzon Strait on the regional atmospheric and weather variations. Long-term multiple satellite observations, including absolute dynamic topography, absolute geostrophic currents, sea surface winds by ASCAT, multi-scale ultra-high resolution sea surface temperature (MURSST) level-four analysis, and research-quality three-hourly TRMM multi-satellite precipitation analysis (TMPA), was used to systematically examine the aforementioned scientific problem. Analysis indicates that the KI is interlinked with the consequential anomalous precipitation off southwestern Taiwan. This anomalous precipitation would lead to ~560 million tons of freshwater influx during each KI event. Subsequently, independent moisture budget analysis suggests that moisture, mainly from vertical advection, is the possible source of the precipitation anomaly. Additionally, a bulk formula analysis was applied to understand how KI can trigger the precipitation anomaly through vertical advection of moisture without causing an evident change in the low-level flows. These new research findings might reconcile the divisiveness on why winds are not showing a synchronous response during the KI and consequential anomalous precipitation events.

The Seasonal Variation of the Anomalously High Salinity at Subsurface Salinity Maximum in Northern South China Sea from Argo Data

The large variations in salinity at the salinity maximum in the northern South China Sea (NSCS), as an indicator for the changes in the Kuroshio intrusion (KI), play an important role in the hydrological cycle. The high salinity here is more than 34.65 at the salinity maximum and is intriguing. In the past, the salinity was difficult to trace in the entire NSCS over long periods due to a lack of high-quality observations. However, due to the availability of accumulated temperature and salinity (T-S) profiles from the Argo program, it is now possible to capture subsurface-maximum data on a large spatiotemporal scale. In this study, the salinity maximum distributed in the subsurface of 80 to 200 m at a density of 23.0–25.5 σθ was extracted from decades of Argo data (on the different pressure surfaces, 2006–2019). We then further studied the spatial distribution and seasonal variation of the salinity maximum and its anomalously high salinity. The results suggest that a high salinity (salinity > 34.65, most of which is located at the shallow depths < 100 m) at the subsurface salinity-maximum layer often occurs in the NSCS, especially near the Luzon Strait, which accounts for about 23% of the total salinity maximum. In winter, the anomalously high salinity at the shallow subsurface salinity maximum can extend to the south of 17° N, while it rarely reaches 18° N and tends to locate at deeper waters in summer. The T-S values of the anomalously high-salinity water are between the mean T-S values in the NSCS and north Pacific subsurface water, implying that the outer sea water gradually mixes with the South China Sea water after passing through the Luzon Strait. Finally, our results show that the factors play an important role in the appearance and distribution of the anomalously high salinity at the subsurface salinity maximum, including the strength of the Kuroshio intrusion, the local wind stress curl and the anticyclonic eddy shedding from the loop current.