The Garlic Tree of Borneo, Scorodocarpus borneensis (Baill.) Becc. (Olacaceae): Potential Utilization in Pharmaceutical, Nutraceutical, and Functional Cosmetic Industries.

Scorodocarpus borneensis (Baill.) Becc. is attracting increased attention as a potential commercial medicinal plant product in Southeast Asia. This review summarizes the current knowledge on the taxonomy, habitat, distribution, medicinal uses, natural products, pharmacology, toxicology, and potential utilization of S. borneesis in the pharmaceutical/nutraceutical/functional cosmetic industries. All data in this review were compiled from Google Scholar, PubMed, Science Direct, Web of Science, ChemSpider, PubChem, and a library search from 1866 to 2022. A total of 33 natural products have been identified, of which 11 were organosulfur compounds. The main organosulfur compound in the seeds is bis-(methylthiomethyl)disulfide, which inhibited the growth of a broad spectrum of bacteria and fungi, T-lymphoblastic leukemia cells, as well as platelet aggregation. Organic extracts evoked anti-microbial, cytotoxic, anti-free radical, and termiticidal effects. S. borneensis and its natural products have important and potentially patentable pharmacological properties. In particular, the seeds have the potential to be used as a source of food preservatives, antiseptics, or termiticides. However, there is a need to establish acute and chronic toxicity, to examine in vivo pharmacological effects and to perform clinical studies.

Ethnobotanical study of medicinal plants used by the Kenyah community of Borneo.

ETHNOPHARMACOLOGICAL RELEVANCE: Sarawak is located in one of the world's most biodiverse regions and is home to more than 40 sub-ethnic groups that each have their own distinct culture, language and lifestyle. This has given rise to numerous, unique ethnobotanical systems. However, due to rapid urbanization, this traditional knowledge is at a risk of extinction. Yet, ethnobotanical studies in Sarawak are almost non-existent, especially among Orang Ulu communities like the Kenyah. AIM OF STUDY: Therefore, this study was conducted to document the ethnomedicinal knowledge of the Kenyah community. The main objectives of this study are: 1) To determine and document the diversity of medicinal plants used by the Kenyah community, 2) To determine whether the availability of modern medicine has affected Kenyah traditional medicine, and 3) To identify plants which have not been previously cited or used for previously unreported medical uses. MATERIALS AND METHODS: We conducted repeated interviews and field surveys at the Asap-Koyan Resettlement Area, Belaga Sarawak. A total of 24 respondents from four Kenyah longhouses were interviewed in this study. Individuals possessing extensive traditional medicinal knowledge were identified via preliminary interviews or by viva voce. Translators were employed to ensure that there was no miscommunication. The results were evaluated based on the plant's total use-reports and number of respondents citing the plant. The data was also evaluated based on use-reports by ailment category. RESULTS: Over 95% of the respondents were 40 years and older (58.21 years old ± 11.21). This was due to the younger members of the community (40 years old and below) admitting that they had almost no knowledge regarding traditional medicine, as they preferred relying on modern medicine. A total of 61 plant species were mentioned by the 24 respondents Seven plants had five or more respondents citing it, which was more than 20% of the respondents. These plants were Piper betle, Homalomena cordata, Senna alata, Annona muricata, Derris elliptica, Blumea balsamifera and Coscinium fenestratum. CONCLUSION: Almost all of the cited plants had been previously recorded to be used in either Ayurvedic, Chinese herbal medicine, Malay traditional medicine or other Asian ethnomedicinal systems. However, there were four highly cited species that were used for treatments that were scarcely reported in past literature. These were piper betle (used by Kenyah to treat fever), Sauropus andrognus (used by Kenyah to treat fever), Derris elliptica (used by Kenyah to treat fever and influenza) and Coscinuim fenestratum (used by Kenyah to treat toxic effects from non-medical substances).

Logged tropical forests have amplified and diverse ecosystem energetics.

Old-growth tropical forests are widely recognized as being immensely important for their biodiversity and high biomass1. Conversely, logged tropical forests are usually characterized as degraded ecosystems2. However, whether logging results in a degradation in ecosystem functions is less clear: shifts in the strength and resilience of key ecosystem processes in large suites of species have rarely been assessed in an ecologically integrated and quantitative framework. Here we adopt an ecosystem energetics lens to gain new insight into the impacts of tropical forest disturbance on a key integrative aspect of ecological function: food pathways and community structure of birds and mammals. We focus on a gradient spanning old-growth and logged forests and oil palm plantations in Borneo. In logged forest there is a 2.5-fold increase in total resource consumption by both birds and mammals compared to that in old-growth forests, probably driven by greater resource accessibility and vegetation palatability. Most principal energetic pathways maintain high species diversity and redundancy, implying maintained resilience. Conversion of logged forest into oil palm plantation results in the collapse of most energetic pathways. Far from being degraded ecosystems, even heavily logged forests can be vibrant and diverse ecosystems with enhanced levels of ecological function.

The fading popularity of a local ecological calendar from Brunei Darussalam, Borneo.

BACKGROUND: Local ecological calendars are ecocultural frameworks that link temporal and spatial scales, contributing to resilience and adaptive management of natural resources and landscapes. They also facilitate management, access and withdrawal of provisioning ecosystem services. In this article, we describe how the ecological calendar of the Kedayan people of Brunei Darussalam links skyscape and biodiversity with sociocultural aspirations to foster adaptive management of landscape, and provide an understanding of the transmission of calendric knowledge in the community. METHODS: In 2018, we collaborated with sixteen purposively sampled knowledge keepers from the Kedayan community of Brunei Darussalam to document the Kedayan local ecological calendar, and develop a calendrical pictogram. Using a structured questionnaire, we then interviewed 107 randomly selected community members, to understand the contemporary relevance and popularity of the Kedayan calendar, and the transmission of calendric knowledge in the community. RESULTS: Our findings reveal that very few respondents (n = 27, 25.3%) are aware of the existence of Kedayan ecological calendar; majority (n = 80, 74.7%) were not aware of its existence. There is no statistically significant correlation between consulting healers, knowledge on appropriate time requisite to consult healers, and awareness and self-professed knowledge of Kedayan calendar. Only 14 (13.1%) of the respondents reported to have received some form of calendric knowledge, while the majority (86.9%; n = 93) never received any calendric knowledge. Only a negligible 1.9% reported to have transmitted calendric knowledge to others indicating a breakdown in transmission of calendric knowledge. CONCLUSION: The calendric pictogram would help the community in revitalizing their calendar. However, the community will have to invest on enhancing transmission of calendric knowledge.

Therapeutic Potential and Nutraceutical Profiling of North Bornean Seaweeds: A Review.

Malaysia has a long coastline surrounded by various islands, including North Borneo, that provide a suitable environment for the growth of diverse species of seaweeds. Some of the important North Bornean seaweed species are Kappaphycus alvarezii, Eucheuma denticulatum, Halymenia durvillaei (Rhodophyta), Caulerpa lentillifera, Caulerpa racemosa (Chlorophyta), Dictyota dichotoma and Sargassum polycystum (Ochrophyta). This review aims to highlight the therapeutic potential of North Bornean seaweeds and their nutraceutical profiling. North Bornean seaweeds have demonstrated anti-inflammatory, antioxidant, antimicrobial, anticancer, cardiovascular protective, neuroprotective, renal protective and hepatic protective potentials. The protective roles of the seaweeds might be due to the presence of a wide variety of nutraceuticals, including phthalic anhydride, 3,4-ethylenedioxythiophene, 2-pentylthiophene, furoic acid (K. alvarezii), eicosapentaenoic acid, palmitoleic acid, fucoxanthin, ß-carotene (E. denticulatum), eucalyptol, oleic acid, dodecanal, pentadecane (H. durvillaei), canthaxanthin, oleic acid, pentadecanoic acid, eicosane (C. lentillifera), pseudoephedrine, palmitic acid, monocaprin (C. racemosa), dictyohydroperoxide, squalene, fucosterol, saringosterol (D. dichotoma), and lutein, neophytadiene, cholest-4-en-3-one and cis-vaccenic acid (S. polycystum). Extensive studies on the seaweed isolates are highly recommended to understand their bioactivity and mechanisms of action, while highlighting their commercialization potential.

The critical role of natural forest as refugium for generalist species in oil palm-dominated landscapes.

In Borneo, oil palm plantations have replaced much of natural resources, where generalist species tend to be the principal beneficiaries, due to the abundant food provided by oil palm plantations. Here, we analyse the distribution of the Asian water monitor lizard (Varanus salvator) population within an oil palm-dominated landscape in the Kinabatangan floodplain, Malaysian Borneo. By using mark-recapture methods we estimated its population size, survival, and growth in forest and plantation habitats. We compared body measurements (i.e. body weight and body length) of individuals living in forest and oil palm habitats as proxy for the population's health status, and used general least squares estimation models to evaluate its response to highly fragmented landscapes in the absence of intensive hunting pressures. Contrary to previous studies, the abundance of lizards was higher in the forest than in oil palm plantations. Recruitment rates were also higher in the forest, suggesting that these areas may function as a source of new individuals into the landscape. While there were no morphometric differences among plantation sites, we found significant differences among forested areas, where larger lizards were found inhabiting forest adjacent to oil palm plantations. Although abundant in food resources, the limited availability of refugia in oil palm plantations may intensify intra-specific encounters and competition, altering the body size distribution in plantation populations, contrary to what happens in the forest. We conclude that large patches of forest, around and within oil palm plantations, are essential for the dynamics of the monitor lizard population in the Kinabatangan floodplain, as well as a potential source of individuals to the landscape. We recommend assessing this effect in other generalist species, as well as the impact on the prey communities, especially to reinforce the establishment of buffer zones and corridors as a conservation strategy within plantations.

Certified "sustainable" palm oil took the place of endangered Bornean and Sumatran large mammals habitat and tropical forests in the last 30 years.

Tropical forests inhabited by endangered orangutans, rhinos, tigers, and elephants in South-east Asia are threatened by deforestation, including oil palm expansion. Certification has been proposed to label sustainable palm oil production. However, from a remotely sensed time-series and imagery analysis (1984-2020), we discovered that most of the currently certified grower supply bases and concessions in Sumatra and Borneo are located in the 1990s large mammals habitat and in areas that were biodiverse tropical forests less than 30 years ago. In light of this dramatic evidence, we suggest that certification schemes claim for the "sustainable" production of palm oil just because they neglect a very recent past of deforestation and habitat degradation.

Carbon declines along tropical forest edges correspond to heterogeneous effects on canopy structure and function.

Nearly 20% of tropical forests are within 100 m of a nonforest edge, a consequence of rapid deforestation for agriculture. Despite widespread conversion, roughly 1.2 billion ha of tropical forest remain, constituting the largest terrestrial component of the global carbon budget. Effects of deforestation on carbon dynamics in remnant forests, and spatial variation in underlying changes in structure and function at the plant scale, remain highly uncertain. Using airborne imaging spectroscopy and light detection and ranging (LiDAR) data, we mapped and quantified changes in forest structure and foliar characteristics along forest/oil palm boundaries in Malaysian Borneo to understand spatial and temporal variation in the influence of edges on aboveground carbon and associated changes in ecosystem structure and function. We uncovered declines in aboveground carbon averaging 22% along edges that extended over 100 m into the forest. Aboveground carbon losses were correlated with significant reductions in canopy height and leaf mass per area and increased foliar phosphorus, three plant traits related to light capture and growth. Carbon declines amplified with edge age. Our results indicate that carbon losses along forest edges can arise from multiple, distinct effects on canopy structure and function that vary with edge age and environmental conditions, pointing to a need for consideration of differences in ecosystem sensitivity when developing land-use and conservation strategies. Our findings reveal that, although edge effects on ecosystem structure and function vary, forests neighboring agricultural plantations are consistently vulnerable to long-lasting negative effects on fundamental ecosystem characteristics controlling primary productivity and carbon storage.

Imaging spectroscopy reveals the effects of topography and logging on the leaf chemistry of tropical forest canopy trees.

Logging, pervasive across the lowland tropics, affects millions of hectares of forest, yet its influence on nutrient cycling remains poorly understood. One hypothesis is that logging influences phosphorus (P) cycling, because this scarce nutrient is removed in extracted timber and eroded soil, leading to shifts in ecosystem functioning and community composition. However, testing this is challenging because P varies within landscapes as a function of geology, topography and climate. Superimposed upon these trends are compositional changes in logged forests, with species with more acquisitive traits, characterized by higher foliar P concentrations, more dominant. It is difficult to resolve these patterns using traditional field approaches alone. Here, we use airborne light detection and ranging-guided hyperspectral imagery to map foliar nutrient (i.e. P, nitrogen [N]) concentrations, calibrated using field measured traits, over 400 km2 of northeastern Borneo, including a landscape-level disturbance gradient spanning old-growth to repeatedly logged forests. The maps reveal that canopy foliar P and N concentrations decrease with elevation. These relationships were not identified using traditional field measurements of leaf and soil nutrients. After controlling for topography, canopy foliar nutrient concentrations were lower in logged forest than in old-growth areas, reflecting decreased nutrient availability. However, foliar nutrient concentrations and specific leaf area were greatest in relatively short patches in logged areas, reflecting a shift in composition to pioneer species with acquisitive traits. N:P ratio increased in logged forest, suggesting reduced soil P availability through disturbance. Through the first landscape scale assessment of how functional leaf traits change in response to logging, we find that differences from old-growth forest become more pronounced as logged forests increase in stature over time, suggesting exacerbated phosphorus limitation as forests recover.

Bactericidal and cytotoxic activity of a diarylheptanoid (etlingerin) isolated from a ginger (Etlingera pubescens) endemic to Borneo.

AIMS: The aim of this study was to investigate the antimicrobial activities of Etlingera pubescens, and to isolate and identify the antimicrobial compound. METHODS AND RESULTS: The crude extracts of E. pubescens were obtained through methanol extraction, and evaluated for antimicrobial activities. From this extract, 1,7-bis(3,4-dihydroxyphenyl)heptan-3-yl acetate (etlingerin) was isolated. When compared to curcumin (a compound with a similar chemical structure), etlingerin showed twofold lower minimum inhibitory concentration values while also being bactericidal. Through time kill assay, etlingerin showed rapid killing effects (as fast as 60 min) against the Gram-positive bacteria (Staphylococcus aureus ATCC 43300 and Bacillus subtilis ATCC 8188). Further assessment revealed that etlingerin caused leakage of intracellular materials, therefore suggesting alteration in membrane permeability as its antimicrobial mechanism. Cytotoxicity study demonstrated that etlingerin exhibited approximately 5- to 12-fold higher IC50 values against several cell lines, as compared to curcumin. CONCLUSIONS: Etlingerin isolated from E. pubescens showed better antibacterial and cytotoxic activities when compared to curcumin. Etlingerin could be safe for human use, though further cytotoxicity study using animal models is needed. SIGNIFICANCE AND IMPACT OF THE STUDY: Etlingerin has a potential to be used in treating bacterial infections due to its good antimicrobial activity, while having potentially low cytotoxicity.

Leucoxenols A and B, two new phenolics from Bornean medicinal plant Syzygium leucoxylon.

The medicinal plant, Syzygium leucoxylon or commonly known as Obah found in North Borneo was considered as traditional medicine by local committee. Two new phenolics, leucoxenols A (1) and B (2) were isolated and identified as major secondary metabolites from the leaves of S. leucoxylon. Their chemical structures were elucidated based on spectroscopic data such as NMR and HRESIMS. Furthermore, these compounds were active against selected strains of fungi.

Phosphorus and nitrogen resorption from different chemical fractions in senescing leaves of tropical tree species on Mount Kinabalu, Borneo.

Nutrient resorption, a process by which plants degrade organic compounds and resorb their nutrients from senescing tissues, is a crucial plant function to increase growth and fitness in nutrient-poor environments. Tropical trees on phosphorus (P)-poor soils are particularly known to have high P-resorption efficiency (PRE, the percentage of P resorbed from senescing leaves before abscission per total P in green leaves). However, the biochemical mechanisms underlying this greater PRE remain unclear. In this study, we determined the P concentration in easily soluble, nucleic acid, lipid and residual fractions for green and senescent leaves of 22 tree species from three sites, which differed in P availability, on the lower flanks of Mt. Kinabalu, Borneo. PRE varied from 24 to 93% and was higher in species from the P-poor site. P-resorption rate was greatest from the lipid fraction, the nucleic acid fraction, and lowest in the easily soluble fraction and the residual fraction when all the species were pooled. For species with higher PRE, P-resorption rate of the residual fraction was relatively high and was comparable in magnitude to that of the other labile fractions. This suggests that tree species inhabiting P-poor environments increased PRE by improving the degradation of recalcitrant compounds. This study suggests that plants selectively degrade organic compounds depending on environmental conditions, which is a key mechanism underlying the variation of PRE.

Over two decades of Plasmodium knowlesi infections in Sarawak: Trend and forecast.

Malaria is still of great public health concern, especially in Malaysian Borneo. The aim of this study was to determine the trends of P. knowlesi infection in Sarawak, Malaysia and to forecast the incidence of P. knowlesi until the year 2040. Data on P. knowlesi malaria cases from 1992 to the year 2014 were obtained from the Sarawak Health Department, Malaysia. ARIMA model was applied to forecast the future incidence of P. knowlesi infection. The data for the whole of Sarawak and subsequently the selected six districts which have high incidence rates of P. knowlesi infection were analyzed. Results of the analysis showed that there was an increasing trend of P. knowlesi cases from the year 1992-2014 (p<0.001). The trend in the incidence started to increase in the year 2008 (p=0.029). The incidence rate per 100,000 populations was between 4.15 in the year 1992 and 42.03 in the year 2014. High incidence of P. knowlesi infections has been detected in the districts adjacent to each other within the interior region of Sarawak. The forecasted incidence and incidence rate per 100,000 populations in the year 2020 were 1229 and 44.04, respectively, while those in the year 2040 were 2056 and 62.91, respectively. The forecasted incidence showed an upward trend highlighting an urgent need to draw up strategic and holistic prevention plans to limit further the increase in P. knowlesi morbidity and mortality in Sarawak. It is imperative that these measures are customized taking into consideration the challenges faced in the interior areas of Sarawak and the behavior of the main vector of P. knowlesi (i.e., An. latens) in Sarawak.

Investigation on the antimicrobial activities of gingers (Etlingera coccinea (Blume) S.Sakai & Nagam and Etlingera sessilanthera R.M.Sm.) endemic to Borneo.

AIMS: To investigate the antimicrobial properties of Etlingera coccinea and Etlingera sessilanthera and to isolate and identify the antimicrobial compounds. METHODS AND RESULTS: Extracts were obtained via sequential solvent extraction method using hexane, dichloromethane, ethyl acetate, methanol and water. Antimicrobial activity testing was done using broth microdilution assay against 17 strains of bacteria. The leaf hexane extract of E. coccinea and rhizome hexane extract of E. sessilanthera showed best antimicrobial activities, with minimum inhibitory concentration (MIC) values ranging from 0·016 to 1 mg ml-1 against Gram-positive bacteria. From these active extracts, two antimicrobials were isolated and identified as trans-2-dodecenal and 8(17),12-labdadiene-15,16-dial with MIC values ranging from 4 to 8 µg ml-1 against Bacillus cereus, Bacillus subtilis and Staphylococcus aureus. CONCLUSION: Etlingera coccinea and E. sessilanthera demonstrated good antimicrobial activities against clinically relevant bacteria strains. The antimicrobial compounds isolated showed low MIC values, hence suggesting their potential use as antimicrobial agents. SIGNIFICANCE AND IMPACT OF THE STUDY: This study is the first to identify the potent antimicrobials from these gingers. The antimicrobials isolated could potentially be developed further for use in treatment of bacterial infections. Also, this study warrants further research into other Etlingera species in search for more antimicrobial compounds.

Capgermacrenes D-G, new sesquiterpenoids from a Bornean soft coral, Capnella imbricata.

Four new bicyclogermacrenes, capgermacrenes D (1) E (2) F (3) and G (4) were isolated from a population of Bornean soft coral Capnella imbricata. The structures of these metabolites were elucidated based on their nuclear magnetic resonance and high-resolution electrospray ionisation mass spectrometry spectral data. These compounds showed bacteriastatic and bacteriacidal activities against Staphylococcus aureus and methicillin-resistant Staphylococcus aureus.

Metabolic profiling and in vitro assessment of anthelmintic fractions of Picria fel-terrae Lour.

Anthelmintic resistance is widespread in gastrointestinal nematode populations, such that there is a consistent need to search for new anthelmintics. However, the cost of screening for new compounds is high and has a very low success rate. Using the knowledge of traditional healers from Borneo Rainforests (Sarawak, Malaysia), we have previously shown that some traditional medicinal plants are a rich source of potential new anthelmintic drug candidates. In this study, Picria fel-terrae Lour. plant extract, which has previously shown promising anthelmintic activities, was fractionated via the use of a solid phase extraction cartridge and each isolated fraction was then tested on free-living nematode Caenorhabditis elegans and the parasitic nematode Haemonchus contortus. We found that a single fraction was enriched for nematocidal activity, killing ≥90% of C. elegans adults and inhibiting the motility of exsheathed L3 of H. contortus, while having minimal cytotoxic activity in mammalian cell culture. Metabolic profiling and chemometric analysis of the effective fraction indicated medium chained fatty acids and phenolic acids were highly represented.

Nutrient allocation among plant organs across 13 tree species in three Bornean rain forests with contrasting nutrient availabilities.

Allocation of nitrogen (N) and phosphorus (P) among plant organs is an important factor regulating growth rate, which is a key ecological process associated with plant life-history strategies. However, few studies have explored how N and P investment in photosynthetic (leaves) and non-photosynthetic (stems and roots) organs changes in relation to depletion of each element. We investigated nutrient concentrations of plant organs in relation to whole-plant nutrient concentration (total nutrient weight per total biomass) as an index of nutrient status of each individual using the saplings of the 13 species in three tropical rain forests with contrasting N and P availabilities (tropical evergreen forests and tropical heath forests). We found a steeper decrease in foliar N concentration than foliar P concentration with decreasing whole-plant nutrient concentration. Moreover, the steeper decrease in foliar N concentration was associated with relatively stable N concentration in stems, and vice versa for P. We suggest that the depletion of N is associated with a rapid dilution of foliar N because the cell walls in non-photosynthetic organs function as an N sink. On the other hand, these species can maintain foliar P concentration by decreasing stem P concentrations despites the depletion of P. Our results emphasize the significance of non-photosynthetic organs as an N sink for understanding the variation of foliar nutrient concentrations for the tree species in the three Bornean rain forests with different N and P availabilities.

Floral diversity and pollination strategies of three rheophytic Schismatoglottideae (Araceae).

Homoplastic evolution of 'unique' morphological characteristics in the Schismatoglottideae - many previously used to define genera - prompted this study to compare morphology and function in connection with pollination biology for Aridarum nicolsonii, Phymatarum borneense and Schottarum sarikeense. Aridarum nicolsonii and P. borneense extrude pollen through a pair of horned thecae while S. sarikeense sheds pollen through a pair of pores on the thecae. Floral traits of spathe constriction, presence and movement of sterile structures on the spadix, the comparable role of horned thecae and thecae pores, the presence of stamen-associated calcium oxalate packages, and the timing of odour emission are discussed in the context of their roles in pollinator management. Pollinators for all investigated species were determined to be species of Colocasiomyia (Diptera: Drosophilidae).

A New Seco-Clerodane-Type Diterpenoid from Bornean Liverwort Schistochila acuminata.

A new compound, schistochilic acid D (1) and two known compounds (2 and 3) were isolated from MeOH extract of Bornean liverwort. Schistochila acuininata collected from Mount Trus Madi, Sabah. The structure of the new metabolite was established based on spectroscopic (ID NMR, 2D NMR, and IR). and HRESIMS data. In addition, another population of S. acuminata collected from Mount Alab (Sabah) yielded four known compounds, 2, 3, 4 and 5. These compounds were tested for their biological potential against the B 16-Fl0 cell line. Compounds 4 and 5 exhibited weak cytotoxic activity.

North Atlantic forcing of tropical Indian Ocean climate.

The response of the tropical climate in the Indian Ocean realm to abrupt climate change events in the North Atlantic Ocean is contentious. Repositioning of the intertropical convergence zone is thought to have been responsible for changes in tropical hydroclimate during North Atlantic cold spells, but the dearth of high-resolution records outside the monsoon realm in the Indian Ocean precludes a full understanding of this remote relationship and its underlying mechanisms. Here we show that slowdowns of the Atlantic meridional overturning circulation during Heinrich stadials and the Younger Dryas stadial affected the tropical Indian Ocean hydroclimate through changes to the Hadley circulation including a southward shift in the rising branch (the intertropical convergence zone) and an overall weakening over the southern Indian Ocean. Our results are based on new, high-resolution sea surface temperature and seawater oxygen isotope records of well-dated sedimentary archives from the tropical eastern Indian Ocean for the past 45,000 years, combined with climate model simulations of Atlantic circulation slowdown under Marine Isotope Stages 2 and 3 boundary conditions. Similar conditions in the east and west of the basin rule out a zonal dipole structure as the dominant forcing of the tropical Indian Ocean hydroclimate of millennial-scale events. Results from our simulations and proxy data suggest dry conditions in the northern Indian Ocean realm and wet and warm conditions in the southern realm during North Atlantic cold spells.