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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: 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., (© 2022 Botanical Society of America.)

Premise: Angiosperm leaves present a classic identification problem due to their morphological complexity. Computer-vision algorithms can identify diagnostic regions in images, and heat map outputs illustrate those regions for identification, providing novel insights through visual feedback. We investigate the potential of analyzing leaf heat maps to reveal novel, human-friendly botanical information with applications for extant- and fossil-leaf identification., Methods: We developed a manual scoring system for hotspot locations on published computer-vision heat maps of cleared leaves that showed diagnostic regions for family identification. Heat maps of 3114 cleared leaves of 930 genera in 14 angiosperm families were analyzed. The top-5 and top-1 hotspot regions of highest diagnostic value were scored for 21 leaf locations. The resulting data were viewed using box plots and analyzed using cluster and principal component analyses. We manually identified similar features in fossil leaves to informally demonstrate potential fossil applications., Results: The method successfully mapped machine strategy using standard botanical language, and distinctive patterns emerged for each family. Hotspots were concentrated on secondary veins (Salicaceae, Myrtaceae, Anacardiaceae), tooth apices (Betulaceae, Rosaceae), and on the little-studied margins of untoothed leaves (Rubiaceae, Annonaceae, Ericaceae). Similar features drove the results from multivariate analyses. The results echo many traditional observations, while also showing that most diagnostic leaf features remain undescribed., Conclusions: Machine-derived heat maps that initially appear to be dominated by noise can be translated into human-interpretable knowledge, highlighting paths forward for botanists and paleobotanists to discover new diagnostic botanical characters., (© 2022 Botanical Society of America.)

The Malay Archipelago is one of the most biodiverse regions on Earth, but it suffers high extinction risks due to severe anthropogenic pressures. Paleobotanical knowledge provides baselines for the conservation of living analogs and improved understanding of vegetation, biogeography, and paleoenvironments through time. The Malesian bioregion is well studied palynologically, but there have been very few investigations of Cenozoic paleobotany (plant macrofossils) in a century or more. We report the first paleobotanical survey of Brunei Darussalam, a sultanate on the north coast of Borneo that still preserves the majority of its extraordinarily diverse, old-growth tropical rainforests. We discovered abundant compression floras dominated by angiosperm leaves at two sites of probable Pliocene age: Berakas Beach, in the Liang Formation, and Kampong Lugu, in an undescribed stratigraphic unit. Both sites also yielded rich palynofloral assemblages from the macrofossil-bearing beds, indicating lowland fern-dominated swamp (Berakas Beach) and mangrove swamp (Kampong Lugu) depositional environments. Fern spores from at least nine families dominate both palynological assemblages, along with abundant fungal and freshwater algal remains, rare marine microplankton, at least four mangrove genera, and a diverse rainforest tree and liana contribution (at least 19 families) with scarce pollen of Dipterocarpaceae, today's dominant regional life form. Compressed leaves and rare reproductive material represent influx to the depocenters from the adjacent coastal rainforests. Although only about 40% of specimens preserve informative details, we can distinguish 23 leaf and two reproductive morphotypes among the two sites. Dipterocarps are by far the most abundant group in both compression assemblages, providing rare, localized evidence for dipterocarp-dominated lowland rainforests in the Malay Archipelago before the Pleistocene. The dipterocarp fossils include winged Shorea fruits, at least two species of plicate Dipterocarpus leaves, and very common Dryobalanops leaves. We attribute additional leaf taxa to Rhamnaceae ( Ziziphus ), Melastomataceae, and Araceae ( Rhaphidophora ), all rare or new fossil records for the region. The dipterocarp leaf dominance contrasts sharply with the family's <1% representation in the palynofloras from the same strata. This result directly demonstrates that dipterocarp pollen is prone to strong taphonomic filtering and underscores the importance of macrofossils for quantifying the timing of the dipterocarps' rise to dominance in the region. Our work shows that complex coastal rainforests dominated by dipterocarps, adjacent to swamps and mangroves and otherwise similar to modern ecosystems, have existed in Borneo for at least 4-5 million years. Our findings add historical impetus for the conservation of these gravely imperiled and extremely biodiverse ecosystems., Competing Interests: Peter Wilf is an Academic Editor for PeerJ. David Shaw is employed by Biostratigraphic Associates (UK) Ltd., and Joseph J. Lambiase is the director of Lambiase Geoscience., (© 2022 Wilf et al.)

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.

Premise: Eocene floras of Patagonia document biotic response to the final separation of Gondwana. The conifer genus Araucaria, distributed worldwide during the Mesozoic, has a disjunct extant distribution between South America and Australasia. Fossils assigned to Australasian Araucaria Sect. Eutacta usually are represented by isolated organs, making diagnosis difficult. Araucaria pichileufensis E.W. Berry, from the middle Eocene Río Pichileufú (RP) site in Argentine Patagonia, was originally placed in Sect. Eutacta and later reported from the early Eocene Laguna del Hunco (LH) locality. However, the relationship of A. pichileufensis to Sect. Eutacta and the conspecificity of the Araucaria material among these Patagonian floras have not been tested using modern methods., Methods: We review the type material of A. pichileufensis alongside large (n = 192) new fossil collections of Araucaria from LH and RP, 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., Results: We describe Araucaria huncoensis sp. nov. from LH and improve the whole-plant concept for Araucaria pichileufensis from RP. The two species respectively resolve in the crown and stem of Sect. Eutacta., Conclusions: Our results confirm the presence and indicate the survival of Sect. Eutacta in South America during early Antarctic separation. The exceptionally complete fossils significantly predate several molecular age estimates for crown Eutacta. The differentiation of two Araucaria species demonstrates conifer turnover during climate change and initial South American isolation from the early to middle Eocene., (© 2020 Botanical Society of America.)

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.)

Evolutionary divergence-age estimates derived from molecular 'clocks' are frequently correlated with paleogeographic, paleoclimatic and extinction events. One prominent hypothesis based on molecular data states that the dominant pattern of Southern Hemisphere biogeography is post-Gondwanan clade origins and subsequent dispersal across the oceans in a metaphoric 'Green Web'. We tested this idea against well-dated Patagonian fossils of 19 plant lineages, representing organisms that actually lived on Gondwana. Most of these occurrences are substantially older than their respective, often post-Gondwanan molecular dates. The Green Web interpretation probably results from directional bias in molecular results. Gondwanan history remains fundamental to understanding Southern Hemisphere plant radiations, and we urge significantly greater caution when using molecular dating to interpret the biological impacts of geological events., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

Unlabelled: •, Premise of Study: Have Gondwanan rainforest floral associations survived? Where do they occur today? Have they survived continuously in particular locations? How significant is their living floristic signal? We revisit these classic questions in light of significant recent increases in relevant paleobotanical data.•, Methods: We traced the extinction and persistence of lineages and associations through the past across four now separated regions-Australia, New Zealand, Patagonia, and Antarctica-using fossil occurrence data from 63 well-dated Gondwanan rainforest sites and 396 constituent taxa. Fossil sites were allocated to four age groups: Cretaceous, Paleocene-Eocene, Neogene plus Oligocene, and Pleistocene. We compared the modern and ancient distributions of lineages represented in the fossil record to see if dissimilarity increased with time. We quantified similarity-dissimilarity of composition and taxonomic structure among fossil assemblages, and between fossil and modern assemblages.•, Key Results: Strong similarities between ancient Patagonia and Australia confirmed shared Gondwanan rainforest history, but more of the lineages persisted in Australia. Samples of ancient Australia grouped with the extant floras of Australia, New Guinea, New Caledonia, Fiji, and Mt. Kinabalu. Decreasing similarity through time among the regional floras of Antarctica, Patagonia, New Zealand, and southern Australia reflects multiple extinction events.•, Conclusions: Gondwanan rainforest lineages contribute significantly to modern rainforest community assembly and often co-occur in widely separated assemblages far from their early fossil records. Understanding how and where lineages from ancient Gondwanan assemblages co-occur today has implications for the conservation of global rainforest vegetation, including in the Old World tropics., (© 2014 Botanical Society of America, Inc.)

Widespread deforestation, agriculture, and construction of milldams by European settlers greatly influenced valley-bottom stream morphology and riparian vegetation in the northeastern USA. The former broad, tussock-sedge wetlands with small, anastomosing channels were converted into today's incised, meandering streams with unstable banks that support mostly weedy, invasive vegetation. Vast accumulations of fine-grained "legacy" sediments that blanket the regional valley-bottom Piedmont landscape now are being reworked from stream banks, significantly impairing the ecological health of downstream water bodies, most notably the Chesapeake Bay. However, potential restoration is impaired by lack of direct knowledge of the pre-settlement riparian and upslope floral ecosystems. We studied the subfossil leaf flora of Denlingers Mill, an obsolete (breached) milldam site in southeastern Pennsylvania that exhibits a modern secondary forest growing atop thin soils, above bedrock outcrops immediately adjacent to a modified, incised stream channel. Presumably, an overhanging old-growth forest also existed on this substrate until the early 1700s and was responsible for depositing exceptionally preserved, minimally transported subfossil leaves into hydric soil strata, which immediately underlie post-European settlement legacy sediments. We interpret the eleven identified species of the subfossil assemblage to primarily represent a previously unknown, upland Red Oak-American Beech mixed hardwood forest. Some elements also appear to belong to a valley-margin Red Maple-Black Ash swamp forest, consistent with preliminary data from a nearby site. Thus, our results add significantly to a more complete understanding of the pre-European settlement landscape, especially of the hardwood tree flora. Compared with the modern forest, it is apparent that both lowland and upslope forests in the region have been modified significantly by historical activities. Our study underscores that generally overlooked subfossil leaves can provide important, local, temporally constrained paleoecological data, with much potential value in this case for stream and wetland restoration decisions in the mid-Atlantic region.

We report the first fossil pollen from South America of the lineage that includes the recently discovered, extremely rare Australian Wollemi Pine, Wollemia nobilis (Araucariaceae). The grains are from the late Paleocene to early middle Eocene Ligorio Márquez Formation of Santa Cruz, Patagonia, Argentina, and are assigned to Dilwynites, the fossil pollen type that closely resembles the pollen of modern Wollemia and some species of its Australasian sister genus, Agathis. Dilwynites was formerly known only from Australia, New Zealand, and East Antarctica. The Patagonian Dilwynites occurs with several taxa of Podocarpaceae and a diverse range of cryptogams and angiosperms, but not Nothofagus. The fossils greatly extend the known geographic range of Dilwynites and provide important new evidence for the Antarctic region as an early Paleogene portal for biotic interchange between Australasia and South 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.

Although temporal calibration is widely recognized as critical for obtaining accurate divergence-time estimates using molecular dating methods, few studies have evaluated the variation resulting from different calibration strategies. Depending on the information available, researchers have often used primary calibrations from the fossil record or secondary calibrations from previous molecular dating studies. In analyses of flowering plants, primary calibration data can be obtained from macro- and mesofossils (e.g., leaves, flowers, and fruits) or microfossils (e.g., pollen). Fossil data can vary substantially in accuracy and precision, presenting a difficult choice when selecting appropriate calibrations. Here, we test the impact of eight plausible calibration scenarios for Nothofagus (Nothofagaceae, Fagales), a plant genus with a particularly rich and well-studied fossil record. To do so, we reviewed the phylogenetic placement and geochronology of 38 fossil taxa of Nothofagus and other Fagales, and we identified minimum age constraints for up to 18 nodes of the phylogeny of Fagales. Molecular dating analyses were conducted for each scenario using maximum likelihood (RAxML + r8s) and Bayesian (BEAST) approaches on sequence data from six regions of the chloroplast and nuclear genomes. Using either ingroup or outgroup constraints, or both, led to similar age estimates, except near strongly influential calibration nodes. Using "early but risky" fossil constraints in addition to "safe but late" constraints, or using assumptions of vicariance instead of fossil constraints, led to older age estimates. In contrast, using secondary calibration points yielded drastically younger age estimates. This empirical study highlights the critical influence of calibration on molecular dating analyses. Even in a best-case situation, with many thoroughly vetted fossils available, substantial uncertainties can remain in the estimates of divergence times. For example, our estimates for the crown group age of Nothofagus varied from 13 to 113 Ma across our full range of calibration scenarios. We suggest that increased background research should be made at all stages of the calibration process to reduce errors wherever possible, from verifying the geochronological data on the fossils to critical reassessment of their phylogenetic position.

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.

Present-day correlations between leaf physiognomic traits (shape and size) and climate are widely used to estimate paleoclimate using fossil floras. For example, leaf-margin analysis estimates paleotemperature using the modern relation of mean annual temperature (MAT) and the site-proportion of untoothed-leaf species (NT). This uniformitarian approach should provide accurate paleoclimate reconstructions under the core assumption that leaf-trait variation principally results from adaptive environmental convergence, and because variation is thus largely independent of phylogeny it should be constant through geologic time. Although much research acknowledges and investigates possible pitfalls in paleoclimate estimation based on leaf physiognomy, the core assumption has never been explicitly tested in a phylogenetic comparative framework. Combining an extant dataset of 21 leaf traits and temperature with a phylogenetic hypothesis for 569 species-site pairs at 17 sites, we found varying amounts of non-random phylogenetic signal in all traits. Phylogenetic vs. standard regressions generally support prevailing ideas that leaf-traits are adaptively responding to temperature, but wider confidence intervals, and shifts in slope and intercept, indicate an overall reduced ability to predict climate precisely due to the non-random phylogenetic signal. Notably, the modern-day relation of proportion of untoothed taxa with mean annual temperature (NT-MAT), central in paleotemperature inference, was greatly modified and reduced, indicating that the modern correlation primarily results from biogeographic history. Importantly, some tooth traits, such as number of teeth, had similar or steeper slopes after taking phylogeny into account, suggesting that leaf teeth display a pattern of exaptive evolution in higher latitudes. This study shows that the assumption of convergence required for precise, quantitative temperature estimates using present-day leaf traits is not supported by empirical evidence, and thus we have very low confidence in previously published, numerical paleotemperature estimates. However, interpreting qualitative changes in paleotemperature remains warranted, given certain conditions such as stratigraphically closely-spaced samples with floristic continuity.

Insect herbivores are considered vulnerable to extinctions of their plant hosts. Previous studies of insect-damaged fossil leaves in the US Western Interior showed major plant and insect herbivore extinction at the Cretaceous-Palaeogene (K-T) boundary. Further, the regional plant-insect system remained depressed or ecologically unbalanced throughout the Palaeocene. Whereas Cretaceous floras had high plant and insect-feeding diversity, all Palaeocene assemblages to date had low richness of plants, insect feeding or both. Here, we use leaf fossils from the middle Palaeocene Menat site, France, which has the oldest well-preserved leaf assemblage from the Palaeocene of Europe, to test the generality of the observed Palaeocene US pattern. Surprisingly, Menat combines high floral diversity with high insect activity, making it the first observation of a 'healthy' Palaeocene plant-insect system. Furthermore, rich and abundant leaf mines across plant species indicate well-developed host specialization. The diversity and complexity of plant-insect interactions at Menat suggest that the net effects of the K-T extinction were less at this greater distance from the Chicxulub, Mexico, impact site. Along with the available data from other regions, our results show that the end-Cretaceous event did not cause a uniform, long-lasting depression of global terrestrial ecosystems. Rather, it gave rise to varying regional patterns of ecological collapse and recovery that appear to have been strongly influenced by distance from the Chicxulub structure.

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.

Food web recovery from mass extinction is poorly understood. We analyzed insect-feeding damage on 14,999 angiosperm leaves from 14 latest Cretaceous, Paleocene, and early Eocene sites in the western interior United States. Most Paleocene floras have low richness of plants and of insect damage. However, a low-diversity 64.4-million-year-old flora from southeastern Montana shows extremely high insect damage richness, especially of leaf mining, whereas an anomalously diverse 63.8-million-year-old flora from the Denver Basin shows little damage and virtually no specialized feeding. These findings reveal severely unbalanced food webs 1 to 2 million years after the end-Cretaceous extinction 65.5 million years ago.

South America has some of the most diverse floras and insect faunas that are known, but its Cenozoic fossil record of insects and insect herbivory is sparse. We quantified insect feeding on 3,599 leaves from the speciose Laguna del Hunco flora (Chubut, Argentina), which dates to the early Eocene climatic optimum (52 million years ago) and compared the results with three well preserved, rich, and identically analyzed early- and middle-Eocene floras from the following sites in North America: Republic, WA; Green River, UT; and Sourdough, WY. We found significantly more damage diversity at Laguna del Hunco than in the North American floras, whether measured on bulk collections or on individual plant species, for both damage morphotypes and feeding groups. An ancient history of rich, specialized plant-insect associations on diverse plant lineages in warm climates may be a major factor contributing to the current biodiversity 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.

Summary: Fossil discoveries can transform our understanding of plant diversification over time and space. Recently described fossils in many plant families have pushed their known records farther back in time, pointing to alternative scenarios for their origin and spread.Here, we describe two new Eocene fossil berries of the nightshade family (Solanaceae) from the Esmeraldas Formation in Colombia and the Green River Formation in Colorado (USA). The placement of the fossils was assessed using clustering and parsimony analyses based on 10 discrete and five continuous characters, which were also scored in 291 extant taxa.The Colombian fossil grouped with members of the tomatillo subtribe, and the Coloradan fossil aligned with the chili pepper tribe. Along with two previously reported early Eocene fossils from the tomatillo genus, these findings indicate that Solanaceae were distributed at least from southern South America to northwestern North America by the early Eocene.Together with two other recently discovered Eocene berries, these fossils demonstrate that the diverse berry clade and, in turn, the entire nightshade family, is much older and was much more widespread in the past than previously thought. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]