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Eocene floras of Patagonia document biotic response to the final separation of Gondwana. The conifer genusAraucaria, distributed worldwide during the Mesozoic, has a disjunct extant distribution between South America and Australasia. Fossils assigned to AustralasianAraucariaSect.Eutactausually are represented by isolated organs, making diagnosis difficult.Araucaria pichileufensisE.W. Berry, from the middle Eocene Río Pichileufú (RP) site in Argentine Patagonia, was originally placed in Sect.Eutactaand later reported from the early Eocene Laguna del Hunco (LH) locality. However, the relationship ofA. pichileufensisto Sect.Eutactaand the conspecificity of theAraucariamaterial among these Patagonian floras have not been tested using modern methods.

We review the type material ofA. pichileufensisalongside large (n= 192) new fossil collections ofAraucariafromLHandRP, including multi‐organ preservation of leafy branches, ovuliferous complexes, and pollen cones. We use a total evidence phylogenetic analysis to analyze relationships of the fossils to Sect.Eutacta.

We describeAraucaria huncoensissp. nov. fromLHand improve the whole‐plant concept forAraucaria pichileufensisfromRP. The two species respectively resolve in the crown and stem of Sect.Eutacta.

Our results confirm the presence and indicate the survival of Sect.Eutactain South America during early Antarctic separation. The exceptionally complete fossils significantly predate several molecular age estimates for crownEutacta. The differentiation of twoAraucariaspecies demonstrates conifer turnover during climate change and initial South American isolation from the early to middle Eocene.

In extant ecosystems, complex networks of ecological interactions between organisms can be readily studied. In contrast, understanding of such interactions in ecosystems of the geologic past is incomplete. Specifically, in past terrestrial ecosystems we know comparatively little about plant biotic interactions besides saprotrophy, herbivory, mycorrhizal associations, and oviposition. Due to taphonomic biases, epiphyte communities are particularly rare in the plant-fossil record, despite their prominence in modern ecosystems. Accordingly, little is known about how terrestrial epiphyte communities have changed across geologic time. Here, we describe a tinyin situfossil epiphyte community that sheds light on plant-animal and plant-plant interactions more than 50 million years ago.

A single silicifiedTodea(Osmundaceae) rhizome from a new locality of the early Eocene (ca. 52 Ma) Tufolitas Laguna del Hunco (Patagonia, Argentina) was studied in serial thin sections using light microscopy. The community of organisms colonizing the tissues of the rhizome was characterized by identifying the organisms and mapping and quantifying their distribution. A 200 × 200 µm grid was superimposed onto the rhizome cross section, and the colonizers present at each node of the grid were tallied.

Preservedin situ, this community offers a rare window onto aspects of ancient ecosystems usually lost to time and taphonomic processes. The community is surprisingly diverse and includes the first fossilized leafy liverworts in South America, also marking the only fossil record of leafy bryophyte epiphytes outside of amber deposits; as well as several types of fungal hyphae and spores; microsclerotia with possible affinities in several ascomycete families; and evidence for oribatid mites.

The community associated with the Patagonian rhizome enriches our understanding of terrestrial epiphyte communities in the distant past and adds to a growing body of literature on osmundaceous rhizomes as important hosts for component communities in ancient ecosystems, just as they are today. Because osmundaceous rhizomes represent an ecological niche that has remained virtually unchanged over time and space and are abundant in the fossil record, they provide a paleoecological model system that could be used to explore epiphyte community structure through time.