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Abstract PremiseAsia's wet tropical forests face a severe biodiversity crisis, but few fossils record their evolutionary history. We recently discovered in situ cuticles on fossil leaves, attributed to the giant rainforest treeDryobalanopsof the iconic Dipterocarpaceae family, from the Plio‐Pleistocene of Brunei Darussalam (northern Borneo). Studying these specimens allowed us to validate the generic identification and delineate affinities to living dipterocarp species. MethodsWe compared the leaf cuticles and architecture of these fossil leaves with the seven livingDryobalanopsspecies. ResultsThe cuticular features shared between the fossils and extantDryobalanops, including the presence of giant stomata on veins, confirm their generic placement. The leaf characters are identical to those ofD. rappa, an IUCN red‐listed Endangered, northern Borneo endemic. TheD. rappamonodominance at the fossil site, along withDipterocarpusspp. leaf fossils, indicates a dipterocarp‐dominated forest near the mangrove‐swamp depocenter, most likely in an adjacent peatland. ConclusionsTheDryobalanops rappafossils are the first fossil evidence of a living endangered tropical tree species and show how analysis of in situ cuticles can help illuminate the poorly known floristic history of the Asian tropics. This discovery highlights new potential for fossils to inform heritage values and paleoconservation in Southeast Asia.more » « lessFree, publicly-accessible full text available May 8, 2026
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Abstract Marine sedimentary rocks of the late Eocene Pagat Member of the Tanjung Formation in the Asem Asem Basin near Satui, Kalimantan, provide an important geological archive for understanding the paleontological evolution of southern Kalimantan (Indonesian Borneo) in the interval leading up the development of the Central Indo-Pacific marine biodiversity hotspot. In this paper, we describe a moderately diverse assemblage of marine invertebrates within a sedimentological and stratigraphical context. In the studied section, the Pagat Member of the Tanjung Formation records an interval of overall marine transgression and chronicles a transition from the marginal marine and continental siliciclastic succession in the underlying Tambak Member to the carbonate platform succession in the overlying Berai Formation. The lower part of the Pagat Member contains heterolithic interbedded siliciclastic sandstone and glauconitic shale, with thin bioclastic floatstone and bioclastic rudstone beds. This segues into a calcareous shale succession with common foraminiferal packstone/rudstone lenses interpreted as low-relief biostromes. A diverse trace fossil assemblage occurs primarily in a muddy/glauconitic sandstone, sandy mudstone, and bioclastic packstone/rudstone succession, constraining the depositional setting to a mid-ramp/mid to distal continental shelf setting below fair-weather wave base but above storm wave base. Each biostrome rests upon a storm-generated ravinement surface characterized by a low-diversityGlossifungitesorTrypanitestrace fossil assemblage. The erosional surfaces were colonized by organisms that preferred stable substrates, including larger benthic foraminifera, solitary corals, oysters, and serpulid annelid worms. The biostromes comprised islands of high marine biodiversity on the mud-dominated Pagat coastline. Together, the biostromes analyzed in this study contained 13 genera of symbiont-bearing larger benthic foraminifera, ~40 mollusk taxa, at least 5 brachyuran decapod genera, and 6 coral genera (Anthemiphyllia,Balanophyllia,Caryophyllia,Cycloseris,Trachyphyllia, andTrochocyathus), as well as a variety of bryozoans, serpulids, echinoids, and asterozoans. High foraminiferal and molluscan diversity, coupled with modest coral diversity, supports the hypothesis that the origin of the diverse tropical invertebrate faunas that characterize the modern Indo-Australian region may have occurred in the latest Eocene/earliest Oligocene.more » « lessFree, publicly-accessible full text available February 1, 2026
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First fossil-leaf floras from Brunei Darussalam show dipterocarp dominance in Borneo by the PlioceneThe Malay Archipelago is one of the most biodiverse regions on Earth, but it suffers high extinction risks due to severe anthropogenic pressures. Paleobotanical knowledge provides baselines for the conservation of living analogs and improved understanding of vegetation, biogeography, and paleoenvironments through time. The Malesian bioregion is well studied palynologically, but there have been very few investigations of Cenozoic paleobotany (plant macrofossils) in a century or more. We report the first paleobotanical survey of Brunei Darussalam, a sultanate on the north coast of Borneo that still preserves the majority of its extraordinarily diverse, old-growth tropical rainforests. We discovered abundant compression floras dominated by angiosperm leaves at two sites of probable Pliocene age: Berakas Beach, in the Liang Formation, and Kampong Lugu, in an undescribed stratigraphic unit. Both sites also yielded rich palynofloral assemblages from the macrofossil-bearing beds, indicating lowland fern-dominated swamp (Berakas Beach) and mangrove swamp (Kampong Lugu) depositional environments. Fern spores from at least nine families dominate both palynological assemblages, along with abundant fungal and freshwater algal remains, rare marine microplankton, at least four mangrove genera, and a diverse rainforest tree and liana contribution (at least 19 families) with scarce pollen of Dipterocarpaceae, today’s dominant regional life form. Compressed leaves and rare reproductive material represent influx to the depocenters from the adjacent coastal rainforests. Although only about 40% of specimens preserve informative details, we can distinguish 23 leaf and two reproductive morphotypes among the two sites. Dipterocarps are by far the most abundant group in both compression assemblages, providing rare, localized evidence for dipterocarp-dominated lowland rainforests in the Malay Archipelago before the Pleistocene. The dipterocarp fossils include winged Shorea fruits, at least two species of plicate Dipterocarpus leaves, and very common Dryobalanops leaves. We attribute additional leaf taxa to Rhamnaceae ( Ziziphus ), Melastomataceae, and Araceae ( Rhaphidophora ), all rare or new fossil records for the region. The dipterocarp leaf dominance contrasts sharply with the family’s <1% representation in the palynofloras from the same strata. This result directly demonstrates that dipterocarp pollen is prone to strong taphonomic filtering and underscores the importance of macrofossils for quantifying the timing of the dipterocarps’ rise to dominance in the region. Our work shows that complex coastal rainforests dominated by dipterocarps, adjacent to swamps and mangroves and otherwise similar to modern ecosystems, have existed in Borneo for at least 4–5 million years. Our findings add historical impetus for the conservation of these gravely imperiled and extremely biodiverse ecosystems.more » « less
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