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PREMISE OF THE STUDY: The clade Bombacoideae + Malvoideae ('Malvatheca group' sensu Baum et al.) in Malvaceae comprises a mostly tropical lineage with derived taxa that now thrive in higher latitudes. The sparse fossil record, especially for Malvoideae, obscures interpretations of past distributions. We describe fossil leaves of Malvoideae from the middle-late Paleocene Cerrejón Formation in Colombia, which contains evidence for the earliest known neotropical rainforest. METHODS: Fifty-six leaf compressions belonging to Malvaceae were collected from the Cerrejón Formation in northern Colombia. Leaf architectural characters were scored and optimized for 81 genera of Malvaceae. Synapomorphic characters and unique character combinations support natural affinities for the fossil leaves. Fossil pollen from the same formation was also assessed. KEY RESULTS: Despite convergence of overall leaf architecture among many Malvaceae, Malvaciphyllum macondicus sp. nov. can be assigned to the clade Eumalvoideae because of distal and proximal bifurcations of the costal secondary and agrophic veins, a synapomorphy for this clade. CONCLUSIONS: The leaf compressions, the oldest fossils for Eumalvoideae, indicate a minimum divergence time of 58-60 Ma, older than existing estimates from molecular analyses of living species. The abundance of eumalvoid leaves and of bombacoid pollen in the midlate Paleocene of Colombia suggests that the Malvatheca group (Malvoideae + Bombacoideae) was already a common element in neotropical forests and does not support an Australasian origin for Eumalvoideae.

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RESUMO

Neotropical rainforests have a very poor fossil record, making hypotheses concerning their origins difficult to evaluate. Nevertheless, some of their most important characteristics can be preserved in the fossil record: high plant diversity, dominance by a distinctive combination of angiosperm families, a preponderance of plant species with large, smooth-margined leaves, and evidence for a high diversity of herbivorous insects. Here, we report on an approximately 58-my-old flora from the Cerrejón Formation of Colombia (paleolatitude approximately 5 degrees N) that is the earliest megafossil record of Neotropical rainforest. The flora has abundant, diverse palms and legumes and similar family composition to extant Neotropical rainforest. Three-quarters of the leaf types are large and entire-margined, indicating rainfall >2,500 mm/year and mean annual temperature >25 degrees C. Despite modern family composition and tropical paleoclimate, the diversity of fossil pollen and leaf samples is 60-80% that of comparable samples from extant and Quaternary Neotropical rainforest from similar climates. Insect feeding damage on Cerrejón fossil leaves, representing primary consumers, is abundant, but also of low diversity, and overwhelmingly made by generalist feeders rather than specialized herbivores. Cerrejón megafossils provide strong evidence that the same Neotropical rainforest families have characterized the biome since the Paleocene, maintaining their importance through climatic phases warmer and cooler than present. The low diversity of both plants and herbivorous insects in this Paleocene Neotropical rainforest may reflect an early stage in the diversification of the lineages that inhabit this biome, and/or a long recovery period from the terminal Cretaceous extinction.

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RESUMO

Palms are a monophyletic group with a dominantly tropical distribution; however, their fossil record in low latitudes is strikingly scarce. In this paper, we describe fossil leaves, inflorescences, and fruits of palms from the middle to late Paleocene Cerrejón Formation, outcropping in the Ranchería River Valley, northern Colombia. The fossils demonstrate the presence of at least five palm morphospecies in the basin ca. 60 Ma. We compare the morphology of the fossils with extant palms and conclude that they belong to at least three palm lineages: the pantropical Cocoseae of the subfamily Arecoideae, the monotypic genus Nypa, and either Calamoideae or Coryphoideae. The fossil fruits and inflorescences are among the oldest megafossil records of these groups and demonstrate that the divergence of the Cocoseae was more than 60 Ma, earlier than has previously been thought. These fossils are useful in tracing the range expansion or contraction of historical or current neotropical elements and also have profound implications for the understanding of the evolution of neotropical rainforests.

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Both the fossil record and molecular data support a long evolutionary history for the Araceae. Although the family is diverse in tropical America today, most araceous fossils, however, have been recorded from middle and high latitudes. Here, we report fossil leaves of Araceae from the middle-late Paleocene of northern Colombia, and review fossil araceous pollen grains from the same interval. Two of the fossil leaf species are placed in the new fossil morphogenus Petrocardium Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov.; these fossils are very similar in leaf morphology to extant Anthurium; however, their relationship to the genus is still unresolved. A third fossil leaf type from Cerrejón is recognized as a species of the extant genus Montrichardia, the first fossil record for this genus. These fossils inhabited a coastal rainforest ∼60-58 million years ago with broadly similar habitat preferences to modern Araceae.

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Tropical South America has the highest plant diversity of any region today, but this richness is usually characterized as a geologically recent development (Neogene or Pleistocene). From caldera-lake beds exposed at Laguna del Hunco in Patagonia, Argentina, paleolatitude approximately 47 degrees S, we report 102 leaf species. Radioisotopic and paleomagnetic analyses indicate that the flora was deposited 52 million years ago, the time of the early Eocene climatic optimum, when tropical plant taxa and warm, equable climates reached middle latitudes of both hemispheres. Adjusted for sample size, observed richness exceeds that of any other Eocene leaf flora, supporting an ancient history of high plant diversity in warm areas of South America.

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