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Leguminosae are one of the most diverse flowering-plant groups today, but the evolutionary history of the family remains obscure because of the scarce early fossil record, particularly from lowland tropics. Here, we report ~500 compression or impression specimens with distinctive legume features collected from the Cerrejón and Bogotá Formations, Middle to Late Paleocene of Colombia. The specimens were segregated into eight fruit and six leaf morphotypes. Two bipinnate leaf morphotypes are confidently placed in the Caesalpinioideae and are the earliest record of this subfamily. Two of the fruit morphotypes are placed in the Detarioideae and Dialioideae. All other fruit and leaf morphotypes show similarities with more than one subfamily or their affinities remain uncertain. The abundant fossil fruits and leaves described here show that Leguminosae was the most important component of the earliest rainforests in northern South America c. 60–58 million years ago. 

Taxonomic resolution is a major challenge in palynology, largely limiting the ecological and evolutionary interpretations possible with deep-time fossil pollen data. We present an approach for fossil pollen analysis that uses optical superresolution microscopy and machine learning to create a quantitative and higher throughput workflow for producing palynological identifications and hypotheses of biological affinity. We developed three convolutional neural network (CNN) classification models: maximum projection (MPM), multislice (MSM), and fused (FM). We trained the models on the pollen of 16 genera of the legume tribe Amherstieae, and then used these models to constrain the biological classifications of 48 fossilStriatopollisspecimens from the Paleocene, Eocene, and Miocene of western Africa and northern South America. All models achieved average accuracies of 83 to 90% in the classification of the extant genera, and the majority of fossil identifications (86%) showed consensus among at least two of the three models. Our fossil identifications support the paleobiogeographic hypothesis that Amherstieae originated in Paleocene Africa and dispersed to South America during the Paleocene-Eocene Thermal Maximum (56 Ma). They also raise the possibility that at least three Amherstieae genera (Crudia,Berlinia, andAnthonotha) may have diverged earlier in the Cenozoic than predicted by molecular phylogenies.

 

While modern forests have their origin in the diversification and expansion of angiosperms in the late Cretaceous and early Cenozoic, it is unclear if the rise of closed-canopy tropical rainforests preceded or followed the end-Cretaceous extinction. The “canopy effect” is a strong vertical gradients in the carbon isotope (δ13C) composition of leaves in modern closed-canopy forests that could serve as a proxy signature for canopy structure in ancient forests. To test this, we report measurements of the carbon isotope composition of nearly 200 fossil angiosperm leaves from two localities in the Paleocene Cerrejón Formation and one locality in the Maastrichtian Guaduas Formation. Leaves from one Cerrejón fossil assemblage deposited in a small fluvial channel exhibited a 6.3‰ range in δ13C, consistent with a closed-canopy forest. Carbon isotope values from lacustrine sediments in the Cerrejón Fm. had a range of 3.3‰, consistent with vegetation along a lake edge. An even narrower range of δ13C values (2.7‰) was observed for a leaf assemblage recovered from the Cretaceous Guaduas Fm., and suggests vegetation with an open canopy structure. Carbon isotope fractionation by late Cretaceous and early Paleogene leaves was in all cases similar to modern relatives, consistent with estimates of low atmospheric CO2 during this time period. This study confirms other lines of evidence suggesting closed-canopy forests in tropical South America existed by the late Paleocene, and fails to find isotopic evidence for a closed-canopy forest in the Cretaceous. 

Leguminosae are one of the most diverse flowering-plant groups today, but the evolutionary history of the family remains obscure because of the scarce early fossil record, particularly from lowland tropics. Here, we report ~500 compression or impression specimens with distinctive legume features collected from the Cerrejón and Bogotá Formations, Middle to Late Paleocene of Colombia. The specimens were segregated into eight fruit and six leaf morphotypes. Two bipinnate leaf morphotypes are confidently placed in the Caesalpinioideae and are the earliest record of this subfamily. Two of the fruit morphotypes are placed in the Detarioideae and Dialioideae. All other fruit and leaf morphotypes show similarities with more than one subfamily or their affinities remain uncertain. The abundant fossil fruits and leaves described here show that Leguminosae was the most important component of the earliest rainforests in northern South America c. 60–58 million years ago.