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The evolutionary history of Eucalyptus and the eucalypts, the larger clade of seven genera including Eucalyptus that today have a natural distribution almost exclusively in Australasia, is poorly documented from the fossil record. Little physical evidence exists bearing on the ancient geographical distributions or morphologies of plants within the clade. Herein, we introduce fossil material of Eucalyptus from the early Eocene (ca. 51.9 Ma) Laguna del Hunco paleoflora of Chubut Province, Argentina; specimens include multiple leaves, infructescences, and dispersed capsules, several flower buds, and a single flower. Morphological similarities that relate the fossils to extant eucalypts include leaf shape, venation, and epidermal oil glands; infructescence structure; valvate capsulate fruits; and operculate flower buds. The presence of a staminophore scar on the fruits links them to Eucalyptus, and the presence of a transverse scar on the flower buds indicates a relationship to Eucalyptus subgenus Symphyomyrtus. Phylogenetic analyses of morphological data alone and combined with aligned sequence data from a prior study including 16 extant eucalypts, one outgroup, and a terminal representing the fossils indicate that the fossils are nested within Eucalyptus. These are the only illustrated Eucalyptus fossils that are definitively Eocene in age, and the only conclusively identified extant or fossil eucalypts naturally occurring outside of Australasia and adjacent Mindanao. Thus, these fossils indicate that the evolution of the eucalypt group is not constrained to a single region. Moreover, they strengthen the taxonomic connections between the Laguna del Hunco paleoflora and extant subtropical and tropical Australasia, one of the three major ecologic-geographic elements of the Laguna del Hunco paleoflora. The age and affinities of the fossils also indicate that Eucalyptus subgenus Symphyomyrtus is older than previously supposed. Paleoecological data indicate that the Patagonian Eucalyptus dominated volcanically disturbed areas adjacent to standing rainforest surrounding an Eocene caldera lake.

Extant neotropical rainforest biomes are characterized by a high diversity and abundance of angiosperm trees and vines, high proportions of entire-margined leaves, high proportions of large leaves (larger than 4500 mm2), high abundance of drip tips and a suite of characteristic dominant families: Sapotaceae, Lauraceae, Leguminosae (Fabaceae), Melastomataceae and Palmae (Arecaceae). Our aim is to define parameters of extant rainforests that will allow their recognition in the fossil record of South America and to evaluate all known South American plant fossil assemblages for first evidence and continued presence of those parameters. We ask when did these critical rainforest characters arise? When did vegetative parameters reach the level of abundance that we see in neotropical forests? Also, when do specific lineages become common in neotropical forests? Our review indicates that evidence of neotropical rainforest is exceedingly rare and equivocal before the Palaeocene. Even in the Palaeocene, the only evidence for tropical rainforest in South America is the appearance of moderately high pollen diversity. By contrast, North American sites provide evidence that rainforest leaf physiognomy was established early in the Palaeocene. By the Eocene in South America, several lines of evidence suggest that neotropical rainforests were diverse, physiognomically recognizable as rainforest and taxonomically allied to modern neotropical rainforests. A mismatch of evidence regarding the age of origin between sites of palaeobotanical high diversity and sites of predicted tropical climates should be reconciled with intensified collecting efforts in South America. We identify several lines of promising research that will help to coalesce previously disparate approaches to the origin, longevity and maintenance of high diversity floras of South America.

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.