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Premise: Two distinct types of fossil infructescences from the early Eocene Laguna del Hunco flora, Chubut Province, Patagonia, Argentina, preserve features of the family Cunoniaceae. The goal of the study was to assess their affinities within Cunoniaceae and to interpret their evolutionary and biogeographical significance. Methods: Specimens were collected from the Tufolitas Laguna del Hunco, Huitrera Formation. They were prepared, photographed, and compared morphologically with similar extant and fossil fruits and infructescences using published literature and herbarium material. Results: The fruit and infructescence morphology place the fossil taxa within Cunoniaceae. They do not conform to any extant genus, supporting the erection of two new fossil genera. Racemofructus gen. nov. shares diagnostic features of the tribe Cunonieae, especially Weinmannia s.l., and exhibits two tribal morphological synapomorphies: a racemose inflorescence and a replum composed of a single column. Cunoniocarpa gen. nov. specimens are paniculate inflorescences with basipetally dehiscent, bicarpellate capsules that have persistent styles and calyces. Its replum morphology suggests an affinity to the tribe Caldcluvieae, particularly to the genus Ackama. Conclusions: The new Patagonian fossils described herein constitute the oldest record of cunoniaceous capsules globally, supplementing a significant body of fossil evidence from pollen, wood, and reproductive structures from southern South America and Antarctica that suggests that the Cunoniaceae were diversified and widely distributed in the southern hemisphere by the early Eocene. Racemofructus and Cunoniocarpa are, respectively, the first fossil records from South America of reproductive structures with affinity to tribes Cunonieae and Caldcluvieae. [ABSTRACT FROM AUTHOR]

Premise of research. The early Eocene Laguna del Hunco (LH) fossil site, northwestern Chubut Province, Argentina, holds one of the best-preserved and most diverse paleofloras worldwide. The paleoflora comprises ferns, gymnosperms, and flowering plants. Despite the rapidly growing knowledge of its macrofossil record, little is known about the site's palynological content. Herein, we present the first dispersed spore-pollen assemblages recovered from LH. Methodology. Palynological samples were collected from seven stratigraphic levels of the Tufolitas de LH (Huitrera Formation), of which six yielded palynomorphs. We determine the botanical affinities of fossil morphotypes and provide systematic descriptions of some taxa. In addition, we establish similarities between LH and other Patagonian Eocene localities, and we compare palynological and prior megafloral records from LH. Pivotal results. We identify 56 spore and pollen species and 28 plant families, of which eight (Cyatheaceae, Schizaeaceae, Polypodiaceae, Asteraceae, Chloranthaceae, Nothofagaceae, Rubiaceae, and Ulmaceae) are reliably reported from the site for the first time. Among other Eocene Patagonian palynofloras, the LH assemblage is similar to Pampa de Jones/Nahuel Huapi Este and Confluencia. Conclusions. The LH spore-pollen assemblages augment the plant fossil record for this significant Eocene locality by incorporating new taxa (e.g., Asteraceae, with one morphotype that represents the oldest record of the family in Patagonia). The new data also reinforce the presence of plant families previously reported from macrofossils, such as Juglandaceae, with pollen grains similar to those of the Engelhardia-Alfaroa group, and Fagaceae (Castaneoideae), complementing the macrofossil record of leaves and reproductive structures. [ABSTRACT FROM AUTHOR]

Huncocladus laubenfelsii gen. et sp. nov. is described from the early Eocene (52 million years old) Laguna del Hunco site in Patagonia, Argentina, on the basis of a compression fossil with cuticle remains. The taxon has several similarities with Phyllocladus , together with characters that are absent in extant Phyllocladus species but are otherwise typical of the enclosing scale-leaved clade. Consequently, Huncocladus is interpreted as a relative of Phyllocladus , possibly belonging to its stem group. This view is supported by a phylogenetic analysis of Podocarpaceae, which recovers Huncocladus as sister to Phyllocladus within the here-termed phyllocladoid clade (Phyllocladus + Huncocladus). Huncocladus laubenfelsii is the first macrofossil record of the phyllocladoid lineage in South America or anywhere in the western hemisphere, vastly extending its historical range and constituting an additional lineage shared between Eocene Patagonia and extant and extinct Australasian and South-east Asian rainforests. The disappearance of phyllocladoids from South America adds to the general extinction pattern described previously for southern hemisphere Podocarpaceae, associated with the family's low drought tolerance in the face of climate change (i.e. aridification). Huncocladus is the oldest record of the phyllocladoids, and it represents a new reference point for temporal calibration and biogeographic inference for the evolution of conifers and Australasian rainforests. [ABSTRACT FROM AUTHOR]

The 51.9 Ma Laguna del Hunco (LH) and 47.5 Ma Río Pichileufd (RP) floras from Patagonia, Argentina are unusually rich, angiosperm-dominated assemblages with living relatives in the low-latitude West Pacific, neotropics, and temperate southern latitudes. The diverse gymnosperms in these floras are important for Gondwanan biogeographic history and paleoclimatic interpretations. "Libocedrus" prechilensis Berry 1938 (Cupressaceae), previously known only from the holotype (RP), a vegetative branch, is revised here based on new material from both localities, including a seed cone attached to a shoot with cuticle (LH). Characters of these fossils are diagnostic of monotypic Papuacedrus (highlands of New Guinea and Moluccas). Living P. papuana is most abundant in cloud forests receiving up to 4 m rainfall annually, whereas Austrocedrus (Libocedrus) chilensis, the basis of comparison when the fossil species was named, inhabits dry, cold steppe margins to mediterranean climates in southern South America. We establish Papuacedrus prechilensis comb. nov., which simultaneously invalidates a southern South American connection for the fossil floras and reveals a link to West Pacific montane rainforests. Combined evidence indicates a biome similar to extant subtropical, or tropical montane, rainforests that persisted for at least 4.4 Myr, linking elevated floral richness to abundant rainfall. [ABSTRACT FROM AUTHOR]

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The Eocene Huitrera Formation of northwestern Patagonia, Argentina, is renowned for its diverse, informative, and outstandingly preserved fossil biotas. In northwest Chubut Province, at the Laguna del Hunco locality, this unit includes one of the most diverse fossil floras known from the Eocene, as well as significant fossil insects and vertebrates. It also includes rich fossil vertebrate faunas at the Laguna Fría and La Barda localities. Previous studies of these important occurrences have provided relatively little sedimentological detail, and radioisotopic age constraints are relatively sparse and in some cases obsolete. Here, we describe five fossiliferous lithofacies deposited in four terrestrial depositional environments: lacustrine basin floor, subaerial pyroclastic plain, vegetated, waterlogged pyroclastic lake margin, and extracaldera incised valley. We also report several new 40Ar/39Ar age determinations. Among these, the uppermost unit of the caldera-forming Ignimbrita Barda Colorada yielded a 40Ar/39Ar age of 52.54 ± 0.17 Ma, ~6 m.y. younger than previous estimates, which demonstrates that deposition of overlying fossiliferous lacustrine strata (previously constrained to older than 52.22 ± 0.22 Ma) must have begun almost immediately on the subsiding ignimbrite surface. A minimum age for Laguna del Hunco fossils is established by an overlying ignimbrite with an age of 49.19 ± 0.24 Ma, confirming that deposition took place during the early Eocene climatic optimum. The Laguna Fría mammalian fauna is younger, constrained between a valley-filling ignimbrite and a capping basalt with 40Ar/39Ar ages of 49.26 ± 0.30 Ma and 43.50 ± 1.14 Ma, respectively. The latter age is ~4 m.y. younger than previously reported. These new ages more precisely define the age range of the Laguna Fría and La Barda faunas, allowing greatly improved understanding of their positions with respect to South American mammal evolution, climate change, and geographic isolation. [ABSTRACT FROM AUTHOR]

We document evidence of endophytic oviposition on fossil compression/impression leaves from the early Eocene Laguna del Hunco and middle Eocene Río Pichileufú floras of Patagonia, Argentina. Based on distinctive morphologies and damage patterns of elongate, ovoid, lens-, or teardrop-shaped scars in the leaves, we assign this insect damage to the ichnogenus Paleoovoidus, consisting of an existing ichnospecies, P. rectus, and two new ichnospecies, P. arcuatum and P. bifurcatus. In P. rectus, the scars are characteristically arranged in linear rows along the midvein; in P. bifurcatus, scars are distributed in double rows along the midvein and parallel to secondary veins; and in P. arcuatuin, scars are deployed in rectilinear and arcuate rows. In some cases, the narrow, angulate end of individual scars bear a darkened region encompassing a circular hole or similar feature indicating ovipositor tissue penetration. A comparison to the structure and surface patten of modern ovipositional damage on dicotyledonous leaves suggests considerable similarity to certain zygopteran Odonata. Specifically, members of the Lestidae probably produced P. rectus and P. bifurcatus, whereas species of Coenagrionidae were responsible for P. arcuatum. Both Patagonian localities represent an elevated diversity of potential fern,gymnosperm, and especially angiosperm hosts, the targets of all observed oviposition. However, we did not detect targeting of particular plant families. Our results indicate behavioral stasis for the three ovipositional patterns for at least 50 million years. Nevertheless, synonymy of these oviposition patterns with mid-Mesozoic ichnosDecies indicates older origins for these distinctive modes of ovinosition [ABSTRACT FROM AUTHOR]

Premise: Acmopyle (Podocarpaceae) comprises two extant species from Oceania that are physiologically restricted to ever-wet rainforests, a confirmed fossil record based on leaf adpressions and cuticles in Australia since the Paleocene, and a few uncertain reports from New Zealand, Antarctica, and South America. We investigated fossil specimens with Acmopyle affinities from the early Eocene Laguna del Hunco site in Patagonia, Argentina., Methods: We studied 42 adpression leafy-shoot fossils and included them in a total evidence phylogenetic analysis., Results: Acmopyle grayae sp. nov. is based on heterophyllous leafy shoots with three distinct leaf types. Among these, bilaterally flattened leaves uniquely preserve subparallel, linear features that we interpret as accessory transfusion tissue (ATT, an extra-venous water-conducting tissue). Some apical morphologies of A. grayae shoots are compatible with the early stages of ovuliferous cone development. Our phylogenetic analysis recovers the new species in a polytomy with the two extant Acmopyle species. We report several types of insect-herbivory damage. We also transfer Acmopyle engelhardti from the middle Eocene Río Pichileufú flora to Dacrycarpus engelhardti comb. nov., Conclusions: We confirm the biogeographically significant presence of the endangered West Pacific genus Acmopyle in Eocene Patagonia. Acmopyle is one of the most drought-intolerant genera in Podocarpaceae, possibly due to the high collapse risk of the ATT, and thus the new fossil species provides physiological evidence for the presence of an ever-wet rainforest environment at Laguna del Hunco during the Early Eocene Climatic Optimum., (© 2023 Botanical Society of America.)

Premise: The spurge family Euphorbiaceae is prominent in tropical rainforests worldwide, particularly in Asia. There is little consensus on the biogeographic origins of the family or its principal lineages. No confirmed spurge macrofossils have come from Gondwana., Methods: We describe the first Gondwanan macrofossils of Euphorbiaceae, represented by two infructescences and associated peltate leaves from the early Eocene (52 Myr ago [Ma]) Laguna del Hunco site in Chubut, Argentina., Results: The infructescences are panicles bearing tiny, pedicellate, spineless capsular fruits with two locules, two axile lenticular seeds, and two unbranched, plumose stigmas. The fossils' character combination only occurs today in some species of the Macaranga-Mallotus clade (MMC; Euphorbiaceae), a widespread Old-World understory group often thought to have tropical Asian origins. The associated leaves are consistent with extant Macaranga., Conclusions: The new fossils are the oldest known for the MMC, demonstrating its Gondwanan history and marking its divergence by at least 52 Ma. This discovery makes an Asian origin of the MMC unlikely because immense oceanic distances separated Asia and South America 52 Ma. The only other MMC reproductive fossils so far known are also from the southern hemisphere (early Miocene, southern New Zealand), far from the Asian tropics. The MMC, along with many other Gondwanan survivors, most likely entered Asia during the Neogene Sahul-Sunda collision. Our discovery adds to a substantial series of well-dated, well-preserved fossils from one undersampled region, Patagonia, that have changed our understanding of plant biogeographic history., (© 2023 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Here we present the first record of a stem-Coracii outside the Holarctic region, found in the early Eocene of Patagonia at the Laguna del Hunco locality. Ueekenkcoracias tambussiae gen. et sp. nov. consists of an incomplete right hind limb that presents the following combination of characters, characteristic of Coracii: relatively short and stout tibiotarsus, poorly developed crista cnemialis cranialis, short and wide tarsometatarsus, with the tuberositas m. tibialis cranialis located medially on the shaft, and curved and stout ungual phalanges. Although the presence of a rounded and conspicuous foramen vasculare distale and the trochlea metatarsi II strongly deflected medially resemble Primobucconidae, a fossil group only found in the Eocene of Europe and North America, our phylogenetic analysis indicates the new taxon is the basalmost known Coracii. The unexpected presence of a stem-Coracii in the Eocene of South America indicates that this clade had a more widespread distribution than previously hypothesized, already extending into the Southern Hemisphere by the early Eocene. Ueekenkcoracias tambussiae represents new evidence of the increasing diversity of stem lineages of birds in the Eocene. The new material provides novel morphological data for understanding the evolutionary origin and radiation of rollers and important data for estimates of the divergence time of the group.

Premise: Solanaceae is a scientifically and economically important angiosperm family with a minimal fossil record and an intriguing early evolutionary history. Here, we report a newly discovered fossil lantern fruit with a suite of features characteristic of Physalideae within Solanaceae. The fossil comes from the early Eocene Laguna del Hunco site (ca. 52 Ma) in Chubut, Argentina, which previously yielded the only other physaloid fruit fossil, Physalis infinemundi., Methods: The fruit morphology and calyx venation pattern of the new fossil were compared with P. infinemundi and extant species of Solanaceae., Results: Physalis hunickenii sp. nov. is clearly distinct from P. infinemundi in its fruiting calyx with wider primary veins, longer and thinner lobes, and especially in its venation pattern with high density, transverse tertiary veins; these features support its placement in a new species. In comparison with extant physaloid genera, the calyx venation pattern and other diagnostic traits reinforce placement of the new fossil, like P. infinemundi, within the tribe Physalideae of Solanaceae., Conclusions: Both species of fossil nightshades from Laguna del Hunco represent crown-group Solanaceae but are older than all prior age estimates of the family. Although at least 20 transoceanic dispersals have been proposed as the driver of range expansion of Solanaceae, the Patagonian fossils push back the diversification of the family to Gondwanan times. Thus, overland dispersal across Gondwana is now a likely scenario for at least some biogeographic patterns, in light of the ancient trans-Antarctic land connections between South America and Australia., (© 2020 Botanical Society of America.)

Many plant genera in the tropical West Pacific are survivors from the paleo-rainforests of Gondwana. For example, the oldest fossils of the Malesian and Australasian conifer Agathis (Araucariaceae) come from the early Paleocene and possibly latest Cretaceous of Patagonia, Argentina (West Gondwana). However, it is unknown whether dependent ecological guilds or lineages of associated insects and fungi persisted on Gondwanan host plants like Agathis through time and space. We report insect-feeding and fungal damage on Patagonian Agathis fossils from four latest Cretaceous to middle Eocene floras spanning ca. 18 Myr and compare it with damage on extant Agathis. Very similar damage was found on fossil and modern Agathis, including blotch mines representing the first known Cretaceous-Paleogene boundary crossing leaf-mine association, external foliage feeding, galls, possible armored scale insect (Diaspididae) covers, and a rust fungus (Pucciniales). The similar suite of damage, unique to fossil and extant Agathis, suggests persistence of ecological guilds and possibly the component communities associated with Agathis since the late Mesozoic, implying host tracking of the genus across major plate movements that led to survival at great distances. The living associations, mostly made by still-unknown culprits, point to previously unrecognized biodiversity and evolutionary history in threatened rainforest ecosystems.

The beech-oak family Fagaceae dominates forests from the northern temperate zone to tropical Asia and Malesia, where it reaches its southern limit. We report early Eocene infructescences of Castanopsis , a diverse and abundant fagaceous genus of Southeast Asia, and co-occurring leaves from the 52-million-year-old Laguna del Hunco flora of southern Argentina. The fossil assemblage notably includes many plant taxa that associate with Castanopsis today. The discovery reveals novel Gondwanan history in Fagaceae and the characteristic tree communities of Southeast Asian lower-montane rainforests. The living diaspora associations persisted through Cenozoic climate change and plate movements as the constituent lineages tracked post-Gondwanan mesic biomes over thousands of kilometers, underscoring their current vulnerability to rapid climate change and habitat loss., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Premise of the Study: The fossil record of Agathis historically has been restricted to Australasia. Recently described fossils from the Eocene of Patagonian Argentina showed a broader distribution than found previously, which is reinforced here with a new early Paleocene Agathis species from Patagonia. No previous phylogenetic analyses have included fossil Agathis species., Methods: We describe macrofossils from Patagonia of Agathis vegetative and reproductive organs from the early Danian, as well as leaves with Agathis affinities from the latest Maastrichtian. A total evidence phylogenetic analysis is performed, including the new Danian species together with other fossil species having agathioid affinities., Key Results: Early Danian Agathis immortalis sp. nov. is the oldest definite occurrence of Agathis and one of the most complete Agathis species in the fossil record. Leafy twigs, leaves, pollen cones, pollen, ovuliferous complexes, and seeds show features that are extremely similar to the living genus. Dilwynites pollen grains, associated today with both Wollemia and Agathis and known since the Turonian, were found in situ within the pollen cones., Conclusions: Agathis was present in Patagonia ca. 2 million years after the K-Pg boundary, and the putative latest Cretaceous fossils suggest that the genus survived the K-Pg extinction. Agathis immortalis sp nov. is recovered in a stem position for the genus, while A. zamunerae (Eocene, Patagonia) is recovered as part of the crown. A Mesozoic divergence for the Araucariaceae crown group, previously challenged by molecular divergence estimates, is supported by the combined phylogenetic analyses including the fossil taxa., (© 2018 Botanical Society of America.)

Premise of the Study: The fossil record is critical for testing biogeographic hypotheses. Menispermaceae (moonseeds) are a widespread family with a rich fossil record and alternative hypotheses related to their origin and diversification. The family is well-represented in Cenozoic deposits of the northern hemisphere, but the record in the southern hemisphere is sparse. Filling in the southern record of moonseeds will improve our ability to evaluate alternative biogeographic hypotheses., Methods: Fossils were collected from the Salamanca (early Paleocene, Danian) and the Huitrera (early Eocene, Ypresian) formations in Chubut Province, Argentina. We photographed them using light microscopy, epifluorescence, and scanning electron microscopy and compared the fossils with similar extant and fossil Menispermaceae using herbarium specimens and published literature., Key Results: We describe fossil leaves and endocarps attributed to Menispermaceae from Argentinean Patagonia. The leaves are identified to the family, and the endocarps are further identified to the tribe Cissampelideae. The Salamancan endocarp is assigned to the extant genus Stephania. These fossils significantly expand the known range of Menispermaceae in South America, and they include the oldest (ca. 64 Ma) unequivocal evidence of the family worldwide., Conclusions: Our findings highlight the importance of West Gondwana in the evolution of Menispermaceae during the Paleogene. Currently, the fossil record does not discern between a Laurasian or Gondwanan origin; however, it does demonstrate that Menispermaceae grew well outside the tropics by the early Paleocene. The endocarps' affinity with Cissampelideae suggests that diversification of the family was well underway by the earliest Paleocene., (© 2018 The Authors. American Journal of Botany is published by Wiley Periodicals, Inc. on behalf of the Botanical Society of America.)

Background and Aims: Early Palaeocene (Danian) plant fossils from Patagonia provide information on the recovery from the end-Cretaceous extinction and Cenozoic floristic change in South America. Actinomorphic flowers with eight to ten perianth parts are described and evaluated in a phylogenetic framework. The goal of this study is to determine the identity of these fossil flowers and to discuss their evolutionary, palaeoecological and biogeographical significance., Methods: More than 100 fossilized flowers were collected from three localities in the Danian Salamanca and Peñas Coloradas Formations in southern Chubut. They were prepared, photographed and compared with similar extant and fossil flowers using published literature and herbarium specimens. Phylogenetic analysis was performed using morphological and molecular data., Key Results: The fossil flowers share some but not all the synapomorphies that characterize the Schizomerieae, a tribe within Cunoniaceae. These features include the shallow floral cup, variable number of perianth parts arranged in two whorls, laciniate petals, anthers with a connective extension, and a superior ovary with free styles. The number of perianth parts is doubled and the in situ pollen is tricolporate, with a surface more like that of other Cunoniaceae outside Schizomerieae, such as Davidsonia or Weinmannia., Conclusions: An extinct genus of crown-group Cunoniaceae is recognized and placed along the stem lineage leading to Schizomerieae. Extant relatives are typical of tropical to southern-temperate rainforests, and these fossils likely indicate a similarly warm and wet temperate palaeoclimate. The oldest reliable occurrences of the family are fossil pollen and wood from the Upper Cretaceous of the Antarctica and Argentina, whereas in Australia the family first occurs in upper Palaeocene deposits. This discovery demonstrates that the family survived the Cretaceous-Palaeogene boundary event in Patagonia and that diversification of extant lineages in the family was under way by the earliest Cenozoic., (© The Author(s) 2018. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Premise of the Study: The flip-leaved podocarp Retrophyllum has a disjunct extant distribution in South American and Australasian tropical rainforests and a Gondwanic fossil record since the Eocene. Evolutionary, biogeographic, and paleoecological insights from previously described fossils are limited because they preserve little foliar variation and no reproductive structures., Methods: We investigated new Retrophyllum material from the terminal Cretaceous Lefipán, the early Eocene Laguna del Hunco, and the early/middle Eocene Río Pichileufú floras of Patagonian Argentina. We also reviewed type material of historical Eocene fossils from southern Chile., Key Results: Cretaceous Retrophyllum superstes sp. nov. is described from a leafy twig, while Eocene R. spiralifolium sp. nov. includes several foliage forms and a peduncle with 13 pollen cones. Both species preserve extensive damage from sap-feeding insects associated with foliar transfusion tissue. The Eocene species exhibits a suite of characters linking it to both Neotropical and West Pacific Retrophyllum, along with several novel features. Retrophyllum araucoensis (Berry) comb. nov. stabilizes the nomenclature for the Chilean fossils., Conclusions: Retrophyllum is considerably older than previously thought and is a survivor of the end-Cretaceous extinction. Much of the characteristic foliar variation and pollen-cone morphology of the genus evolved by the early Eocene. The mixed biogeographic signal of R. spiralifolium supports vicariance and represents a rare Neotropical connection for terminal-Gondwanan Patagonia, which is predominantly linked to extant Australasian floras due to South American extinctions. The leaf morphology of the fossils suggests significant drought vulnerability as in living Retrophyllum, indicating humid paleoenvironments., (© 2017 Botanical Society of America.)

Premise of the Study: Eocene caldera-lake beds at Laguna del Hunco (LH, ca. 52.2 Ma) and Río Pichileufú (RP, ca. 47.7 Ma) in Argentine Patagonia provide copious information about the biological history of Gondwana. Several plant genera from these sites are known as fossils from southern Australia and New Zealand and survive only in Australasian rainforests. The potential presence of Dacrycarpus (Podocarpaceae) holds considerable interest due to its extensive foliage-fossil record in Gondwana, its remarkably broad modern distribution in Southeast Asian and Australasian rainforests, its high physiological moisture requirements, and its bird-dispersed seeds. However, the unique seed cones that firmly diagnose Dacrycarpus were not previously known from the fossil record., Methods: I describe and interpret fertile (LH) and vegetative (LH and RP) material of Dacrycarpus and present a nomenclatural revision for fossil Dacrycarpus from South America., Key Results: Dacrycarpus puertae sp. nov. is the first fossil occurrence of the unusual seed cones that typify living Dacrycarpus, attached to characteristic foliage, and of attached Dacrycarpus pollen cones and foliage. Dacrycarpus puertae is indistinguishable from living D. imbricatus (montane, Burma to Fiji). Dacrycarpus chilensis (Engelhardt) comb. nov. is proposed for Eocene vegetative material from Chile., Conclusions: Modern-aspect Dacrycarpus was present in Eocene Patagonia, demonstrating an astonishingly wide-ranging paleogeographic history and implying a long evolutionary association with bird dispersers. Dacrycarpus puertae provides the first significant Asian link for Eocene Patagonian floras, strengthens the biogeographic connections from Patagonia to Australasia across Antarctica during the warm Eocene, and indicates high-rainfall paleoenvironments.

The origins of South America's exceptional plant diversity are poorly known from the fossil record. We report on unbiased quantitative collections of fossil floras from Laguna del Hunco (LH) and Río Pichileufú (RP) in Patagonia, Argentina. These sites represent a frost-free humid biome in South American middle latitudes of the globally warm Eocene. At LH, from 4,303 identified specimens, we recognize 186 species of plant organs and 152 species of leaves. Adjusted for sample size, the LH flora is more diverse than comparable Eocene floras known from other continents. The RP flora shares several taxa with LH and appears to be as rich, although sampling is preliminary. The two floras were previously considered coeval. However, (40)Ar/(39)Ar dating of three ash-fall tuff beds in close stratigraphic association with the RP flora indicates an age of 47.46+/-0.05 Ma, 4.5 million years younger than LH, for which one tuff is reanalyzed here as 51.91+/-0.22 Ma. Thus, diverse floral associations in Patagonia evolved by the Eocene, possibly in response to global warming, and were persistent and areally extensive. This suggests extraordinary richness at low latitudes via the latitudinal diversity gradient, corroborated by published palynological data from the Eocene of Colombia.

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.