Abstract
INTRODUCTION
The Equatorial Pacific Ocean, extending from the Peru–Chile coast to the Indonesian archipelago, has a dominant influence on global ocean–atmosphere interactions. It hosts several tightly linked phenomena, including oscillations in thermocline depth, the Western Pacific Warm Pool (WPWP), El Niño Southern Oscillation (ENSO) variability, the Walker circulation, and changes in high-latitude overturning circulation. These processes collectively shape the global climate by modulating heat distribution, ocean productivity, and atmospheric CO2 exchange. The Eastern Equatorial Pacific (EEP) is particularly important for understanding these linkages due to its strong upwelling, high nutrient concentrations, and persistently shallow thermocline (Wyrtki, 1966). The region is highly sensitive to both tropical and extratropical forcing, making it a key site for studying feedback between ocean circulation and climate variability (Farrell et al., 1995). Palaeoceanographic records (Bin Shaari et al., 2013; Etourneau et al., 2013; Pallone et al., 2025) from the EEP provide valuable insights into how equatorial dynamics have evolved over glacial interglacial cycles and influenced global climate through changes in ocean circulation and biogeochemical cycle, influencing the variation in productivity during the Quaternary. Calcite export fluxes during the Last Glacial Maximum (LGM) were notably lower than during the Holocene, likely reflecting reduced nutrient supply from the Southern Ocean (Loubere et al., 2004). Boron isotope based reconstructions suggest that deep-ocean pH increased during glacial periods, indicating shifts in carbonate chemistry and carbon storage, whereas surface pH remained relatively stable (Sanyal et al., 1997). Despite high macronutrient concentrations, productivity in the EEP is often limited by iron availability, underscoring the importance of nutrient dynamics and ocean–atmosphere carbon exchange in this region (Wells et al., 1999). Although the large scale physical and biogeochemical processes governing EEP variability are increasingly well understood, the detailed faunal and ecological responses to these changes remain less constrained. Understanding these relationships is crucial for interpreting how ocean structure, upwelling strength, and frontal dynamics have evolved over time.
This study examines Quaternary palaeoceanographic variability in the EEP, with a focus on the impact of the Southern Polar Front (SPF). By analysing planktic foraminiferal assemblages, species composition, and abundance patterns from ODP Hole 846B (Figure 1), we aim to reconstruct past surface and thermocline conditions and assess how equatorial ocean dynamics responded to glacial interglacial forcing.
Modern annual Sea Surface Temperature (°C) from the World Ocean Atlas 2023 (Reagan et al., 2024), visualised using Ocean Data View (Schlitzer, 2023) with ODP site 846B (black dot).
MODERN OCEANOGRAPHIC SETUP
In the trans-tropical Pacific, oceanic circulation is largely influenced by tropical atmospheric dynamics and changes in the sign of the Coriolis force across the equator. A defining feature of this circulation is the asymmetry of surface currents on either side of the equator, which corresponds to the mean latitudinal position of the Intertropical Convergence Zone (ITCZ), the region where the northeast and southeast trade winds converge. Along the equator, wind driven divergence depresses the sea surface topography, generating a pressure gradient that drives a westward geostrophic flow on both sides of the equator, known as the South Equatorial Current (SEC). The primary source waters for the SEC are derived from the Peru Current (PC), which transports cold, nutrient rich waters from higher latitudes, and from the eastward flowing subsurface Equatorial Undercurrent (EUC), which contributes to surface waters through upwelling (Mayer et al., 1992).
Site 846B (3°5.696′S, 90°49.078′W) was drilled during Ocean Drilling Programme (ODP) Leg 138 at a water depth of 3,307 m in the EEP. The site lies approximately 300 km south of the Galápagos Islands, within a zone where the SEC interacts with the PC (Figure 2).
Location of ODP Site 846B and present-day oceanographic setup of the Eastern Equatorial Pacific Ocean. Surface current shown as solid arrows; subsurface current as dashed arrows. CAC (California Current); NEC (North Equatorial Current); NECC (North Equatorial Counter current); EUC (Equatorial Undercurrent); SEC (South Equatorial Current); PC (Peru Current). Shaded areas illustrate the general latitudinal extent of the SEC and NEC. (Adopted and modified from Mayer, L., Pisias, N., Janecek, T., et al., 1992 Proceedings of the Ocean Drilling Program, Initial Reports , Vol. 138).
At present, the EEP ‘cold tongue’ functions as a major ocean to atmosphere CO2 source (Takahashi et al., 2009), sustained by the persistent upwelling of cool, nutrient rich subsurface waters supplied primarily by the EUC (Fiedler & Talley, 2006; Kessler, 2006; Toggweiler et al., 1991). These waters originate largely in the Southern Ocean’s Subantarctic Zone, linking high latitude nutrient cycling to low latitude productivity (Rodgers et al., 2003; Toggweiler et al., 1991). Despite elevated macronutrient concentrations, primary productivity in the EEP is constrained by iron and silicic acid limitation, resulting in relatively low phytoplankton biomass even under high nutrient supply (Brzezinski et al., 2011; Dugdale et al., 1995; Quirós-Collazos et al., 2020). The site 846 lies on the southern flank of the Carnegie Ridge, underlain by oceanic crust generated at the Galápagos Spreading Centre. Its palaeopositions have been quantitatively constrained using Pacific–Nazca plate rotation models in conjunction with Pacific hotspot reference frames. Paleogeographic reconstructions indicate that Site 846 has remained within a narrow latitudinal band near its present position throughout much of its geological history (Farrell et al., 1995). This stability makes it a key site for evaluating the temporal evolution of equatorial current systems and large scale ocean circulation patterns.
MATERIALS AND METHODS
The studied core reveals alternating layers of carbonate-rich and silica-rich sediments. Deep sea samples were collected from a 90 meter long core at 30 centimeter intervals. Each sample was processed using the wet sieving method following international standard protocols. Planktic foraminifera retained on the ≥150 µm sieve were studied following Peeters et al. (1999). The samples were weighed and stored in labelled tubes. These specimens were then split to obtain aliquots containing at least 300 individuals, following the methods of Imbrie and Kipp (1971) and Imbrie et al. (1973), and were mounted on assemblage slides. The foraminiferal species were classified based on their water mass and depth habitat preferences within the ocean water column. Taxonomic identification to the species level followed the frameworks of Kennett and Srinivasan (1983), Bolli and Saunders (1985), and Schiebel and Hemleben (2017).
Age Model
Sediment accumulation rates in pelagic settings are primarily controlled by carbonate productivity (Berger & Winterer, 2009). Shackleton et al. (1992) and the Shipboard Scientific Party (Mayer et al., 1992) developed a consistent method to calculate sedimentation rates across multiple drill sites from ODP Leg 138, using a unified stratigraphic framework based on Gamma Ray Attenuation Porosity Evaluator (GRAPE) density measurements. Key stratigraphic events were selected for their close association with well-dated magnetic reversals and biostratigraphic datums, ensuring accurate age control. At ODP Site 846B, sediment accumulation has remained relatively uniform throughout the Quaternary (Figure 3). In the present study, an age model was constructed using established phylogenetic knowledge to assess the reliability of planktonic foraminiferal events.
Nearly uniform sediment accumulation rate at ODP site 846 B.
RESULTS
In this study, 266 ODP samples were analysed. The assemblage is dominated by planktonic foraminiferal species such as
For palaeoceanographic reconstruction during the Quaternary, the planktonic foraminifera were categorised by ecological habitat into three groups: Mixed Layer Dwellers (MLD), dominated by
Relative abundance of Neogloboquadrina dutertrei (Thermocline Dweller) and Globorotalia (Globoconella) inflata (Temperate Group)
According to Loubere (2001), in the EEP,
Relative abundance of N. dutertrei and temperate fauna in sediments from ODP Site 846B. The left panel shows the percentage relative abundance of N. dutertrei (black line) with fluctuations between ~20% and 90%. In contrast, the right panel shows the relative abundance of temperate fauna, dominated by G. inflata during four distinct intervals correlating with estimated glacial periods at 2.1 Ma, 1.2 Ma, 1.0 Ma, and the Late Holocene. Age model and stratigraphic calibration follow Gradstein et al. (2020). This pattern indicates episodic cold-water incursions, likely linked to glacial intensifications in the Southern Hemisphere.
Stratified oceanic systems frequently develop a Deep Chlorophyll Maximum (DCM) that remains undetectable by ocean colour satellites. The DCM may correspond to either an actual phytoplankton biomass peak, termed the Deep Biomass Maximum (DBM), or a physiological adaptation to low-light conditions, in which phytoplankton increase their chlorophyll
During the Early Pleistocene (2.6–1.0 Ma),
The overall pattern reflects a long-term cooling trend and increasing climate variability from the Late Pliocene into the Pleistocene. The dominance of
Periods of increased
The anomalous occurrences of
Variations in relative abundance over time highlight the dominance of
Relative abundance of Mixed Layer Dwellers and Thermocline Dwellers
The MLD group, primarily composed of
Relative abundance of Mixed Layer Dwellers (MLD, left panel) and Thermocline Dwellers (TD, right panel) recorded in ODP Site 846B sediment core. Corresponding age (Ma) is shown. Notable intervals of elevated MLD abundance coincide with declines in TD, indicating dynamic stratification and El Niño like conditions during the Quaternary. Stratigraphic context follows Gradstein et al. (2020). Mixed Layer Census data taken from Mallick et al. (2017).
Six distinct intervals were observed at approximately 2.15 Ma, 2.0 Ma, 1.86 Ma, 1.47 Ma, 1.44 Ma, and 0.57 Ma, showing remarkable increases in the relative abundance of MLD, accompanied by corresponding declines in TD (Figure 5). This inverse relationship suggests El Niño like conditions in the EEP. The species interrelationships and their stratigraphic patterns support this interpretation.
Around 0.5 Ma, MLD reached a peak of ~25.9%, followed by a gradual decline into the Late Pleistocene and Holocene. In contrast, in the youngest part of the core, TD reached its highest relative abundance (~35%), suggesting significant reorganisation of the upper water column during this time.
The observed inverse relationship between MLD and TD abundances reflects alternating phases of surface water stratification and thermocline structure in the EEP. Periods dominated by MLD imply enhanced surface water productivity and a deepened thermocline, conditions often associated with El Niño like states. Conversely, intervals of increased TD abundance indicate a shoaling thermocline, characteristic of intensified upwelling and La Niña like conditions. These long-term oscillations correspond to major climatic transitions through the Pleistocene, including shifts in global ice volume and oceanic circulation that modulated the strength and depth of the EEP thermocline. The parallel trends between species groups and stratigraphic intervals therefore record a dynamic interplay between tropical surface water variability and global climate forcing.
Relative abundance of Upwelling Indicator Species (UIS), Mixed Layer Dwellers (MLD), and Neogloboquadrina dutertrei
The figure compares the relative abundances (percentage) of UIS, MLD, and
Relative abundance profiles of Upwelling Indicator Species (UIS), Mixed layer Dweller and N. dutertrei from ODP Site 846B. and corresponding age (Ma). Major UIS peaks are synchronous with reduced MLD abundance, indicating amplified upwelling and nutrient delivery. Mixed Layer Census data taken from Mallick et al. (2017).
Peaks in UIS percentage are interpreted as episodes of intensified upwelling, likely linked to strengthened trade winds or reorganisations in oceanographic circulation, possibly influenced by major climatic transitions. The marked decrease in MLD during these upwelling intervals indicates increased environmental stress and nutrient enrichment in surface waters, favouring more opportunistic, upwelling adapted taxa. The consistently high abundance of
The comparison thus provides key evidence for the variability and drivers of upwelling in the eastern equatorial during the Quaternary, linking palaeoproductivity changes to global and regional oceanographic controls. The three intervals of maximum UI correspond to major palaeoceanographic reorganisations and can be related to both global climate shifts and changes in regional ocean–atmosphere dynamics.
DISCUSSION
Palaeoceanographic variability in the Eastern Equatorial Pacific
Analysis of planktonic foraminiferal assemblages from ODP Hole 846B documents a dynamic palaeoceanographic evolution of the EEP throughout the Quaternary. The record reveals pronounced fluctuations in water-column structure, upwelling intensity, and intermittent incursions of temperate water masses. When viewed alongside regional and global climate reconstructions, these results highlight the EEP’s sensitivity to both high latitude polar forcing and low latitude atmosphere–ocean interactions.
The EUC serves as the principal source of upwelled water in the eastern Pacific cold tongue (Wyrtki & Kilonsky, 1984). Cromwell (1953) demonstrated that equatorial upwelling arises from the divergence of Ekman transport near the equator. This upwelled water mass, enriched in iron from sedimentary sources along the Papua New Guinea shelf (Ryan et al., 2006), sustains high primary productivity and contributes to deep-ocean CO2 sequestration. Lateral advection from the upper thermocline (Wyrtki, 1981) is consistent with the high relative abundance of
Incursions of temperate fauna and high latitude Linkages
A notable feature of the record is the occurrence of four intervals with anomalously high abundances (≥20%) of the temperate species
The main surface flow paths, indicated by a solid, thick blue arrow, illustrate the intensified Humboldt Current, which has resulted from the northward migration of the Southern Polar Front driven by the expansion of the Antarctic Ice Sheet (AIS). This phenomenon facilitated the unusual invasion of temperate fauna, G. inflata , at the studied ODP 846B at 3˚S latitude. Principal surface flow paths are: PCC (Peru Chile Current), Peru Coastal Current (PCoastalC), Chile Coastal Current (CCoastalC), Peru Chile Counter Current (PCCC), Equatorial Undercurrent (EUC), modified after Karstensen & Ulloa, 2009.
Comparable high latitude linkages have been documented elsewhere in the Southern Hemisphere. Singh and Sinha (2021) identified three major northward migrations of the APF in the southeast Indian Ocean, including one that occurred around 1.2 million years ago (Ma), which coincides with the present event. They associated the proliferation of the
In the EEP, these temperate incursions likely reflect enhanced activity of the Peru–Chile (Humboldt) Current, which advects cold, subpolar water northward (Figure 7). During cooler periods such as the LGM, intensification of this current extended the range of subpolar species along the eastern boundary (Feldberg & Mix, 2003). The synchronous occurrence of temperate fauna in both the Indian and Pacific Oceans suggests a hemisphere-wide oceanographic response to glacial expansion, characterised by stronger eastern boundary currents and increased inter-ocean connectivity (Singh & Sinha, 2021; Sinha & Singh, 2007a, 2007b).
Decoupling of upwelling dynamics
During glacial intervals, particularly the LGM, the EEP experienced pronounced sea-surface cooling, with SSTs decreasing by 5°C–7°C along the equator (Feldberg & Mix, 2003). This was accompanied by a steeper meridional SST gradient south of the equatorial front (Martínez et al., 2003). In the present record, this period is characterised by thermocline shoaling, evidenced by a 35% peak in thermocline dwelling foraminifera, and intensified trade winds, reflected by the dominance of
The intervals of high relative abundance of N. dutertrei at the studied site are due to persistent nutrient supply from the EUC, along with advected water from the Peruvian Margin Upwelling system. Principal surface flow paths are: PCC (Peru Chile Current), Peru Coastal Current (PCoastalC), Chile Coastal Current (CCoastalC), Peru Chile Counter Current (PCCC). Equatorial Undercurrent (EUC), modified after Karstensen & Ulloa, 2009.
The foraminiferal assemblages indicate that two partially decoupled systems maintained EEP upwelling:
Coastal upwelling along the Peruvian margin, dominated by opportunistic taxa such as Equatorial upwelling, driven by trade-wind-induced divergence and sustained by nutrient rich waters of the EUC, is represented chiefly by
Periods of enhanced coastal upwelling did not always coincide with peaks in
Feldberg and Mix (2003) similarly distinguished between high latitude advective and low latitude upwelling components of SST variability. The faunal decoupling observed here provides independent evidence for these distinct yet interacting drivers of regional productivity. This dynamic interplay may explain why the EEP maintained high productivity even during warm phases, challenging simplified models of tropical ocean dynamics.
Water column structure and climatic implications
Variations in the relative abundances of MLD and TD indicate significant reorganisation of upper ocean stratification through time. Six major intervals approximately 2.15 Ma, 2.0 Ma, 1.86 Ma, 1.47 Ma, 1.44 Ma, and 0.5 Ma—show sharp increases in MLD coupled with declines in TD, interpreted as El Niño like conditions with a deepened thermocline and reduced vertical stratification. Conversely, the TD maximum around 27 kyr suggests thermocline shoaling and stronger upwelling, consistent with La Niña like states.
A long-term decline in MLD abundance after 0.5 million years (Ma), accompanied by an increase in TD, signifies a fundamental shift in the EEP ocean structure. This transition may reflect weakening of the trade winds and reduction of the equatorial cold tongue, coinciding with the Mid-Pleistocene Transition (MPT), when glacial interglacial cycles became more intense and climatic variability amplified globally.
CONCLUSIONS
By integrating detailed foraminiferal assemblage data with broader palaeoceanographic evidence, this study provides a high-resolution biological perspective that refines large-scale physical and geochemical interpretations of the EEP during the Quaternary. The record from ODP Hole 846B demonstrates that the EEP was far from static; it experienced significant reorganisations in water-column structure, upwelling strength, and faunal composition driven by both regional and global climate processes.
The repeated presence of
The dominance of
Variations in mixed layer and TD record oscillations between El Niño– and La Niña like conditions, indicating shifts in upper ocean stratification and thermocline depth.
The faunal reorganisation following the MPT signifies a major restructuring of EEP circulation and upwelling systems in response to intensified glacial interglacial cycles.
Overall, the EEP emerges as a climatically sensitive region shaped by an intricate interplay of high-latitude processes, tropical atmospheric dynamics, and boundary current variability.
Footnotes
Acknowledgement
KRM, PKN, and BM acknowledge the infrastructural support provided by the Department of Geology. AKS and DKS express their gratitude to both the Delhi School of Climate Change and Sustainability and the Department of Geology, University of Delhi, for their infrastructural assistance. The research problem was conceptualised by KRM and AKS, who also prepared the manuscript. DKS contributed to drafting the manuscript and provided valuable feedback. Planktic foraminiferal census analyses were performed by PKN and KRM, with technical support from BM.
Declaration of Conflicting Interests
The authors declared no potential conflicts of interest with respect to the research, authorship and/or publication of this article.
Funding
The authors disclosed receipt of the following financial support for the research, authorship and/or publication of this article: The authors gratefully acknowledge funding support provided under the Palaeoclimate Programme of the Ministry of Earth Sciences, Government of India, for carrying out the research, as well as for the preparation and publication of this manuscript (Sanction No. MoES/CCR/Paleo-4/2019).
